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VOL. 113, PARTS 1 & 2 
31 MAY, 1989 

Transactions of the 

Royal Society of South 



Lee, D. C & Birchby, C M. Decoribatula gen. nov. from Singapore, with notes on allied 
Reticuloppia (Acarida: Cryptostigmata: Oribatulidae) from Tropical 
Australia ----------- \ 

Hirst, D. A new genus of huntsman spider (Heteropodidae: Araneae) from south eastern 

Australia - 7 

Tyler, M. J. A new species of Lechriodus (Anura: Leptodactylidae) from the Tertiary of 

Queensland, with a redefinition of the ilial characteristics of the 

genus -- 15 

Boulton, A. J. Over-summering refuges of aquatic macroinvertebrates in two intermittent 

streams in central Victoria 23 

Southcott, R. V. The larva and nymph instars of Odontacarus (Leogonius) adelaideae 

(Womersley) (Acarina: Trombiculidae: Leeuwenhoekiinae) 35 

Tiver, E, Sparrow, A. D. & Lange, R. T. The composition and distribution of the vegetation 

of north-west Eyre Peninsula -------- 47 

Bourman, R. P. & Lindsay, J. M. Timing, extent and character of Late Cainzoic faulting 

on the eastern margin of the Mt Lofty Ranges, South Australia - - 63 

Ainslie, R. G, Johnston, D. A. & Offler, E. W. Intertidal communities of northern Spencer 

Gulf, South Australia - 69 

Koste, W. & Shiel, R. J. Rotifera from Australian inland waters. III. Euchlanidae, 

Mytilinidae and Trichotriidae (Rotifera: Monogononta) 85 

Brief Communications; 

Davies, M. Rediscovery of Uperoleia orientalis (Parker) (Anura: Myobatrachinae) - - 115 

Nobbs, J. The occurrence of plant parasitic nematodes in the arid region of South 

Australia ------ 117 






VOL. 113, PART 1 


CONTENTS, VOL. 113, 1989 

PARTS 1 & 2, 31 May 

Lee, D. C & Birehby, C M. Decoribatula gen. nov. from Singapore, with notes on allied 
Reticuloppia (Acarida: Cryptostigmata: Oribatulidae) from Tropical 
Australia -----..--.. i 

Hirst, D. A new genus of huntsman spider (Heteropodidae: Araneae) from south eastern 

Australia ----------- 7 

Tyler, M. J. A new species of Lechriodus (Anura: Leptodactylidae) from the Tertiary of 
Queensland, with a redefinition of the ilial characteristics of the 
genus - - 15 

Boulton, A. J. Over-summering refuges of aquatic macroinvertebrates in two intermittent 

streams in central Victoria - - 23 

Southcolt, R. V. The larva and nymph instars of Odontacarus (Leogonius) adetaicleae 

(Womersley) (Acarina: Trombieulidae: Leeuwenhoekiinae) - - 35 

Tiver, F., Sparrow, A. D. & Lange, R. T. The composition and distribution of the vegetation 

of north-west Eyre Peninsula ------- 47 

Bourman, R. P. & Lindsay, J. M. Timing, extent and character of Late Cain/.oic faulting 

on the eastern margin of the Mt Lofty Ranges, South Australia - 63 

Ainslie, U. G, Johnston, D. A. & Offler, E. W. Intertidal communities of northern Spencer 

Gulf, South Australia .... - 69 

Koste, W, & Shiel, R. J. Rotifera from Australian inland waters. III. Euchlanidae, 

Mytilinidae and Trichotriidae (Rotifera: Monogononta) - - 85 

Brief Communications: 

Davies, M. Rediscovery of Uperoleia orientalis (Parker) (Anura: Myobatrachinae) - 115 

Nobbs, J. The occurrence of plant parasitic nematodes in the arid region of South 

Australia - - - - 117 

PARTS 3 & 4, 30 November 

Koste, W. & Shiel, R. J. Rotifera from Australian inland waters IV. Colurellidae (Rotifera: 

Monogononta) «- -r *» - 119 

Shorl, A. D., Buckley, R. G, & Fotheringham, D. G. Preliminary investigations of beach 

ridge progradation on Eyre Peninsula and Kangaroo Island - - 145 

Pledge, N. S. The occurrence of a diprotodontid marsupial referable to Euowenia grata 

De Vis in western New South Wales - - - - - -163 

Austin, A. D. & Allen, G. R. Parasitoids of Uraba tugens Walker (Lepidoptera:Noctuidae) 
in South Australia, with description of two new species of 
Braconidae - - — 169 

Barker, S. Contributions to the taxonomy of Australian Buprestidae (Coleoptera): New 
species of As traeus and Stigmodera (Castiarina) and a key to Astraeus 
(s.s) - 185 

Jenkins, J. F. & Hasenohr, P. Trilobites and their trails in a Black Shale: Early Cambrian 

of The Fleurieu Peninsula, South Australia - 195 

Gowlett-Holmcs, K. L. & Holmes, N. J. C Rediscovery of Primovula (P) heleneae Cate 
and description of a new species of Crenavolva from South Australia 
(Mollusca: Gastropoda: Ovulidae) ------ 205 

Davies, M. Developmental biology of the Australopapuan hylid frog Litoria eucnemis 

(Anura: Hylidae) - - - - - - - - - 215 

Brief Communication: 

Beardsell, G. R. Hybridisation of Litoria chloris and L. xanthomera. (Anura: 

Hylidae) - ------- 221 

/nscn to [run sua ions of she Royal Soct<--fy of South A •isimlia, Vot, II?-, parts $ & 4 K 30 November, 1989 




by David C. Lee & Carolyn M. Birchby* 


Decoribatula pustulata gen. nov., sp. nov. on an orchid from Singapore, intercepted at Adelaide 
Airport, is described. It is similar to Reticuloppia reticulata Balogh & Mahunka, 1966 from 
rainforest litter in Queensland, the description of which is extended. Both mites are unusual in 
having a deficient chaetotaxy on femora I and 1 1 . 

KEY WORDS: Acarida, Oribatulidae, Decoribatula pustulata, new genus, new species, 
Reticuloppia reticulata Balogh & Mahunka, Singapore, Queensland, leg chaetotaxy. 



by David C. Leu & Carolyn M. Birchby* 


Ls£, t>. C & Birchey. CM, (1989) Devortba/uta gen. nov. from Singapore, with notes on allied Retivtihppia 
(Aearida: Cryptostigmata: Oribaiuiidae) from tropical Australia. Trans. R. Sov. S Aunt. 113 (2). 1-5, 
31 Ma>; J 989. 

Decorihutufa pusutlota gen. nov., sp. nov. on an orchid from Singapore, intercepted at Adelaide Airport, 
is described. It In similar to Retuuhppia reticulata Balogh & Mahunka, 1966 from rainforest Niter in 
Queensland, the description Of which ffi extended. Both miles are unusual in having a deficient chaetotaxy 
on femora I and II. 

KEY Wokos Avaiida, Onhatulidae, Deconhotula pustulata, new genus, new species, 
Reticulopp'ta reticulata Balogh $ Mahunka, Singapore, Queensland, leg chaetotaxy. 


The mites- considered here have been studied 
because of (heir relevance to an ongoing study of 
saxcoptiform mites of South Australian soils, 
sampled from nine florally diverse sites- An 
introduction to relevant work on the advanced 
an bate mites (Planofissurac), with comments on 
the Oripodoidea, which include the Oribatulidae, 
is provided by Lee <I987). These mites and a group 
of seven species of Oribatula-Vike mites from the 
South Australian study are atypical within the 
Oripodoidea in that they have a deficient chaetotaxy 
on femora I and I). Oripodoid leg chaetotaxy will 
be discussed more fully when the South Australian 
species arc described (Lee & Birchby in preparation). 
The chaelotaxy for femur I and II on oripodoid 
mites is anterior; 2 dorsal/2 ventral, I posterior 
setae (0,2/2,1), In the deficient chaetotaxy one or 
two setae are missing, resulting in one of three 
chactotaxies as follows: 0,2/2,0; 0,2/1,0; 0,2/1,1. The 
Oribfftula-ilke S. Aust mites differ superficially in 
having only short or medium length hysteronotal 
setae and they were collected from only the arid, 
semiarid or malice sites; whilst the two species 
considered here have long hystcronotal setae and 
are from moist, tropical sites. 

The notation and methods of measurements 
follow Lee (1981) with modifications made by Lee 
{1987). Measurements are in micrometers (/tin). The 
type of the new species is deposited in the South 
Australian Museum, Adelaide (SAMA) whilst rhe 
type of Reticufoppia reticulata is deposited in to the 
Hungarian National Museum, Budapest, 

• South Australian Museum, North Terrace, Adelaide. 
S. Aust. 5000 

Hettculoppla Balogh & Mahunka 
ReticutopptD Balo^li d!t Mahunka, 1966; p, 564 

Type-species Reticufoppia reticulata Balogh & 
Mahunka, l%6. 

Diagnosis: Oribauilinae. Hysteronotum with 14 
pairs (5/ f 6Z, 35") of long setae. Lamella absent. 
Hysveronotal lenticulus present near dorsosejugaJ 
furrow. Integument of hysteronotum with extensive- 
reticulate sculpturing and cerotegument forming a 
thick, columnar refractile exudate. Femora 1 and 
U with deficient chactotaxy, lacking posterior setae: 
I - 0,2/2,0; II - 0,2/1,0. Legs slim and long, with 
leg IV longest {leg III is lost). Pretarsa! claws short 
(central claw II less than 0.33 x length of tarsus II) 
and lateral claws much slimmer than central claw 
(depth less than 0.5 x depth of central claw II), 
Remarks' Reticuloppia is based on a single female, 
grouped in the Oribatulinae, and distinguished by 
the hysteronotum having a large number of long 
setae and reticulate sculpturing around a lenticular 
(Balogh & Balogh 1984). In addition, Ihc 
proteronotum lacks ridges, having only a weak 
sublamella, and the leg chactotaxy is deficient in 
a way (hat is unique amongst established oripodoid 
species t 

Reticufoppia reticulata Balogh & Mahunka 

Rcucutoppio reticulata Balogh & Mahunka, 1966; p. 564. 

Female: Idio&omal length, 475 (470 in original 
description). Leg lengths (femur-tarsus); I - 270, 
n - 254, IK-?, TV -311, Tibial maximum heights; 
I - 25,5, II - 20.5. Hi - ?. IV- 18. Yellow brown 
colour. Thick (Maximum depth equal to distance 
between setae d-/2), whitish, columnar retractile 


Figs 1-2: Posterior aspects of femur-pretarsus of right legs I and II, all setae on femora illustrated, p = posterior, 
= ventral. ( s Retkuloppia reticulata Balogh & Mahunka. 2, Decoribatuta pustutata sp. nov. 

cerotegument on proteronotum anterior to ./2, 
thickest lateral to seta zl\ also covering 
hysteronotnm except around lenttculus and near 
pleural margins, with thick patches posterolateral 
to seta 51, lateral to £3 and between setae .76-76. 
Proteronotum with faint cosrate sublamclla. 
Sensory seta (z2) with long stalk (length = x 3 
caput). Margin of hoihridium raised into turret-like 
structure, height subequal to length of ^2 caput. 
Seta a2 short (0.5 x or less distance between setae 
./2-zl), fine, smooth, without cilia. Hysteronoturn 
with fine reticulate sculpturing under columnar 
exudate. Five multiporose foramina* foramen F3 
divided in F5a and Fib. Hysteronotal seta SI 

confirmed similar to and nearly as long as Zl. 
Podosternal chactotaxy: 3/, MI, VII, VV. with all 
third rank setae as long as /V/3 (illustrated in the 
wrong position by Balogh & Mahunka 1966, fig. 
15) and 13 on pedotectum I, If 13 posterior to 
pedotectum II (as for Decoribatuta pusfutosa sp, 
nov. s Fig.4), IV3 on discidium. Seta Sai shorter than 
Sa2 and Sa3. Opisthosternum with foveolatc 
sculpturing. Four subcylindrical eggs, exochorion 
smooth, 80 x 230 (mean). Legs long (mean femur- 
tarsus: 59% of soma) and slim (mean maximum 
tibial height: 27% of mean length). 
Materia/ examined: Holotype female, rainforest 
litter, Mt Spec flifftSttS, I46°1I T E), Townsville, Qld, 

tnxOKMAlUI '..<> Ml res 

I9.iii.l%5, J. Balogh. Some lost, no 
femur-tarsus III on either Mile 
Remarks: The further description here of 
Hctkuinppia reticulata adds lo ur corrects the 
original description, with a lew important 
confirmations. Reference to the relative dimensions 
of the legs is distorted by legs III being lost (usually 
a short, slim teg amongst the oripodoids.i. 

Decoribaiulii gen. nov. 
Type species: Decorihatula pusiulatn sp. nov. 

tyiagnosis: Oribatuliuae. Hysleronotum with 14 
pairs (57, 6Z, 3SS Of mainly long setae (SI is short). 
Lamella with only coslatc anterior part present, not 
reaching bothridium lo seta z2 t Smooth, paler mid- 
dorsal anterior area on hysleronotum, bui not 
clearly delineated as lenticulus. Integument of 
hystcionolum with extensive lovcolalo sculpturing, 
cerotegument inconspicuous. Femora I and II with 
deficient chactotaxy, but with posterior setae: I - 
0,2/1,1; 11-0,2/1,1. Leg* oF medium girth and long, 
with leg IV longest. Prctarsal claws long (centra! 
daw II more than 033 > length of tarsus II) and 
lateral claws only slightly slimmer than central elaw 
(depth more than 0.5 x depth of central claw II), 
Remarks: Decorihatula is based on a single male. 
It is similar to Reikuloppia in having a deficient 
femoral chactotaxy, long hysteronotal setae, a 
divided anterior hysteronotal foramen {FSu, rlh) 
and a turret-like bothridium to c2. It particularly 
differs, in having a lamella, a different type of 
cerolegument, a different type of setal loss on 
femora 1 and II and unusually large pretarsal claws. 
On the basis of this it is considered to be a sister- 
group to Retkuloppta bur different -enough to be 
regarded as a separate genus. 

Deeortbatula pustufata sp. nov. 
FIGS 2-4 
Mole; Idiosomal length, 527. Leg lengths {fctnur- 
taisus): 1-2% 11-280, III -285, IV -32?. Tibial 
maximum heights: I - 36, II - 31, 111 - 23, IV - 
23. Red brown colour, Inconspicuous (depth less 
than diameter of hysteronotal setal bases* 
cernteguirieni. concentrated in round or uval 
pustules on hysteronotum, 

Proteronotum with partial costatc lamella, 
superficially inconspicuous but backed by rclraelilc 
internal apodeme, not reaching, back to turret-like 
bothridium to seta 52. Two multiporose 

foramina <f1. Fid) recognisable Uorsally. Central 
setae <J\sjX z\) with 3 or 4 files of cilia, ,/2 longest. 
Sensory .seta (z2) with caput and exposed stalk 
subequal in length, Seta 52 short (0.6 k or less 
disfancey2-^l), fine, smooth, without cilia. 

Hysteronotum with smooth central triangulate 
anea from anterior margin to level with seta 73, but 
no clearly delineated pale lenticulus. I ive 
multiporose foramina {F'Sa, F3b, F4, F5, F6) each 
backed by a disc-shaped chamber in iniegunienr 
Hysteronotal setae smooth, S\ short, subequai in 
52, whilst 75 and 55 very long with whip-like end. 

Podosternum with reticulaie sculpturing. 
Subpodal ridge in three parts, pedoteetum II and 
diseidiurn large, extending laterally beyond level ol 
pedoteetum I. Lateral setae (especially UB) longer 
than central setae. Opisthosternurn with three setae 
(J/g4 t JZgS t Sg) represented only by bases on both 
sides and assumed broken off. Scia Sa\ lunger than 
Sal and 5a3. Shieid dark, centrally horizontal and 
punctaic with curving upwards, smooth marginal 

Legs long (mean femur-tarsus Strn of soma}, 
with leg I second longest, and ol' medium-girlh 
(mean maximum libial height: 35% of mean 
length), Ventral flange on trochanters III and IV. 
Femora with ventral incrustation. 
Material examined: Holotype male (SAM A 
N1988474), on orchid from Singapore, intercepted 
at Adelaide International Airport, 13.iv. 1987, (free 
Raker (S. Ausi. Department of Agriculture). 
Remarks: The male of D. pustulata is described, 
although nripodoid species are usually based on 
females, because it is unlikely that more material 
will be collected and this species is important lo the 
classification ot'S. Aust. mites being studied (I ee 
& Birchby in preparation). There is a lack of sexual 
dimorphism amongst oripodoid character stale* 
that are considered important in distinguishing 
specjes. The female is likely to be bigger and have 
the thickening around the genital orifice separated 
from the ventrosejugal apodeme. Its relationships 
are considered under the remarks on Decorihatula. 


We thank the Australian Biological Resources 
Study for a grant to D.C.L. funding the salary of 
C.M.B., Ms Kathy Eowshall for the notation and 
presentation of the figures and Mrs Debbie Lowcry 
for typitig the manuscript. 



Fig. 3: Decoribatula pustulaia sp. nov., notum of soma. 



Fig. 4: Decoribatula pustulata sp. nov, sternum of soma, 


Dalogh, J. & Balogh, P. (1984) Review of the 

Oribatuloidea Thor, 1929 (Acari: Oribatei). Acta zool. 

hung. 30, 257-313. 
& Mahunka, S. (1966) New Oribatids (Acari) 

from Australian Soils. Folia ent. hung. 33, 553-568. 
Lee, D. C. (1981) Sarcoptiformes (Acari) of South 

Australian soils. I. Notation. 2. Bifemorata and 

Ptyctima (Cryptostigmata). Rec. S. AusU Mus. 18, 
(1987) Introductory study of advanced oribate 

mites (Acarida: Cryptostigmata: Planofissurae) and a 
redescription of the only valid species of 
Constrictobates (Oripodoidca). Ibid. 21, 35-42. 



by David Hirst* 


A new heteropodine genus, Keilira, gen. nov. and two new species, K sparsornaculata sp. nov. and 
K sokoli sp. nov. are described from South Australia and Victoria respectively. Keilira differs from 
other Australian genera in labium shape and the relatively short, thick embolus of the male. The 
arrangement of cheliceral teeth is similar to that of Heteropoda Latreille, 1804, but the general 
appearance resembles Australian species attributed to Olios Walckenaer, 1837, here considered as 
belonging in Neosparassus Hogg, 1903. 


by David Hirst* 


Hrws» f l>. (1^89) A new genus of huntsman spider (Heteropodidae Aranene) fiom south eastern Austratu 
8i«W /?. Sac. $ AwJ, Hi. 7-/5, 11 May, 19H9. 

A new heteropodine genus, Keilira, yen. nov. and two new species, XI ipamomaadata sp. nov. and 
A*, sokuli sp. nov. are described from Solum Australia and Victoria respectively. Kei/iru differs iron* other 
Australian genera in labium shape and the relatively short, true)* embolus ot the male. The arrangement 
ol chelieeral teeth is similtn to (hat of Ht'tpntpudv I ufreille, 1804, but the general appearance resembles 
Australian species aitt United lo Olios Walckcnaer, 1S?7, hjjpt considered as belonging itt Neosparassus Hogg. 

Key Words. Atancae, Heteropodidae, Ketfiw eon nov., new taxa. 


This, paper forms the first purt of a laxonnmie 
study on I he Australian Heteropodidae. A new 
heteropodine genus is defined lor undeseribed 
^tccies previously mis-identified as Neosparasw 
(subfamily Deteninae Hogg, 1903), which they 
resemble in general colouring and shape. 
Ncosparassus will be discussed fully later (Hirst in 
prep.), but it may be mentioned here that despite 
its revalidation by Jarvi (1914) all relevant species 
h;ivebeen referred to Olios in subsequent literature. 

Subfamilies of the Heteropodidae are poorly 
defined. Some useful characters of the 
Heteropodinne Keyserling, 1H80 are referred to here 
in associating the new genus with that subfamily, 
Similarly; characters o( the Dclcninac are mentioned 
in comparing Neosparassus with Olios, All hough 
limits of thai subfamily arc uncertain, it is retained 
here as separate from the Gusparassinae (Jarvi 1912) 
on I he basis of genitalie structure. 

Materials and Methods 

All measurements are in millimetres, made with 
an eyepiece graticule Eye diameters, interspaces and 
MOQ measuiements arc expressed as relative to the 
diameter of an AME. Drawings were done with the 
aid of a camera tucida on a Wild microscope. All 
abbreviations are standard for the Arancac. Bilateral 
variability oi' leg spine counts are indicated in 
parenthesis. Types are deposited in the South 
Australian Museum, Adelaide (SAM A), and rhe 
Australian Museum, Sydney (AM). 

Ki ji'j'.-j £eiv n«v. 
Diagrtosts; Labium twice as broad as long. Male 
embolus short, thick relatively straight with a curved 

South Austrian Museum, North Terrace, Adelaide, 
S.A*m. 5000, 

tip. Female with small fossa and hood-like anterior 

Defminort: Carapace raised, flatttsh or convex in 
profile, highest in the fovea region, sloping genii y 
to clypcu.v Fovea! groove weak or indistinct. Eyes 
subequal. Anterior eye row slightly recurved. 
Posterior eye row straight lo slightly recurved. 
Clypeus half width of an AMU or slightly less. 
Chelieeral fang groove with three teeth on 
promargin, three or four leeth on retromargin and 
one or more median rows of minute, pointed teeth. 
Labium at least twice as broad as long. Sternum 
barely longer than broad or equal, slightly convex. 
I egs 214.1. Anterior legs with lour ventral pans of 
spines on tibia, without an extreme distal pair 
Reduced piolateral leg spinatiou, patellae without 
prolateraJ spines. Female palp tarsal claw with four 
small teeth. Female genitalia with paired uncoiled 
spermatheeae. Small fossa with weakly selerotued 
hood-like anterior margin. Male embolus short, 
thick, relatively straight with a curved tip. Palpal 
tibia with a small prodorsal distal extension and a 
triangular spur rising from a retrolateral apophysis. 
type species; Keilira sparsomacuhda sp. nov. 


Although resembling \'eosparasms in body shape 
and coloration, the presence of a distal prodorsal 
extension on the tibia of the male palp and three 
teeth on the promargin of the chelicerae followed 
by one oi more row.s of median teeth, places the 
genus close to Heteropodu and Pandercetes L. 
Koch, 1875, the Australian representatives of the 
Heteiopodinac. Neosparassus and other genera of 
the Deleuinae lack median teeth and a well defined 
extension on the male palp tibia. 

The retromarginal teeth of Keilira differ Irorn 
those of Heteropodu and Pandercetes in that both 
the latter have lour long teeth which are evenly sized 
and close together wifh a small space between ihe 
two middle teeth. In Keilira greater spacing occurs 



between the middle *nd distal teeth (rig. 8), This 
placement remains unchanged when a fourth tooth 
iv present, this kiting minute and found al the 
proximal end of the row (Fig. 12>. 

The teeth of the female palp tarsal i-'aws of 
Netervpoda and Pattdercetes are elongated and 
curved, as long or longer than the curved claw tip, 
and comb-like. This condition is approached in at 
least one Other Australian heieropodid, Olios 
hetrtutis Hogg, 1914, the correct generic status of 
which is unknown as no mature specimens, have 
been seen The elongated teeth on I he palpal daw 
;ue nui found in Keiiira, but as aU other Australian 
genera normally possess at least five teeth, it is 
unusual in having four 

Keiiira and Pandereeres have tour ventral spine 
pain on anterior tibiae hut lack a distal pair. While 
four ventia! spine pairs are found in Heteropoda, 
one pair U shouer and positioned distally near the 
articulation with the metatarsi, This character vw 
used by Kaston (J980) to distinguish Hetero/wda 
from Olios, the latter havine two pairs and lackine 
ar- extreme distal pair. NeospamsMts and most other 
Deleninae genera have three pairs with the distal 
pair near the articulation. The exceptions are some 
species of Pediana Simon. 18X0 and juvenile 
specimens of at least teopeda L. Koch, 1875 and 
Neasparassus ijuvenilcs of other genera have not 
been examined). There, the distal spine is 
represented as a bristle barely distinguishable from 
those adjacent do the tibia margin. 

Keiiira is similar to Heteropoila in carapace 
profile and in having leg 1 shorter than leg II 
{Pandweies has a concave carapace profile and kg 
I longer ihan teg II), but diftets in eye position and 
relative size of the anterior median eyev- The mate 
bulb, in the unexpnnded slate, resemble* that of 
Patuiereetes and, to a lesser extenr, Hetempoda, 
Keiiira lacks the long tapering embolus typical of 
other Australian heteropodid genera, having a 
relatively straight, shon, thick embolus which tapers 
at the lip to a blunt point. For much of itv length 
the embolus runs adjacent to the bulb. In 
Panderxwes and Heieropoda the embolus is well 
clear of the bulb for [tt entire length. 

It can be concluded that Keiiira is referable to 
the Heteropodinae, yet is distinct from He?erc>poda 
and Pandereetes, However* it is known lhai 
Heteropodinae genera of uncertain status occur in 
Queensland with live or six vcniral spine pairs on 
anterior tibia. These arc not congeneric with Keiiira 
lunpubl. diita) but appear lo be closely related and 
may prove to be a sister group. 

Keiiira contains I he smallest known Australian 
heteropodids. Its representation by a handful of 
specimens from the south-eastern parts of both 
South Australia and Victoria extends the known 
range of the Australian Heteropodinae. Little is 
known of their biology bul they appear lo have 
similar habits to Neosparassus (Main 1976). Several 
specimens ol Keiiira sparsotnaeulata were collected 
whilst foraging at night on low flowering shrubs 
(LeptQSpeWUtn Junipennum Sm.), which were 
attracting abundant insects, a habitat three specie* 
of Neosparassus have also been found sharing. One 
juvenile specimen of Keiiira was found in litter at 
the base of shrubs during the day. 
Etymology; The generic name Keiiira is derived 
from Keiiira Station, 31 km N.E. of Kingston, S.E, 
South Australia. 

KeiHra tpsrsomacufata sp. nov, 
FIGS 1-10 

Hnlalvpe: SAMA N19S75, & 13 km N. Of 
Keiiira Station. 36* 37* tx 140° IV E, S. Aust„ Feb. 
1974. D. Hirst. 

Allotype: SAMA NI9R76, •? , same data as 

Paratopes; SAMA NI9877, s?, same <3a(a as 
holotvpe; SAMA N 19378, 9, Cortina Station, 
Coorong, 36" 20' S, 139° 46' E, S. Aiisl 20.U96*. 
AAV. l-ornes. 

Other material examined; SAMA N 19872, 
Juvenile, Cortina Station, same data as above; 
SAMA M9873, Juvenile, Kendall Station, Coonong, 
S, Ausl-> Mar. 1968. AAV. Forbes; SAMA NI9874. 
Juvenile, Aldinga Scrub, S> Aust., 16. \x. 1987. D. 

Dcxvnpiion of huhtype (Figs l-4> Carapace length 
2.90, width 2.85. Abdomen length 3.40, width 2.20. 

Colour in alcohol: \ellow-hrown carapace with 
pair of median black, short linear markings with 
adjacent spots, anterior c\' fovea. Caput with darker 
suffusion in ocular region, eyes with black rims. 
Chehccrac rcddiih-ycllow. Sternum yellowish. Legs; 
anterior pairs yellow-orange, posterior pairs yellow, 
palps yellow. Abdomen creamish-white with small 
scattered spots of black pigment, fewer ventTally 

Carapace: Convex, highest between II coxae 
sloping gradualJy Dp clypeus, steeply ro posterior. 
Fovea indistinct, marked by darker pigment. Setae 
sparse short tea 0,2), moderately long (ca 0.4) 
bristles around ocular region, few on caput between 
eyes and fovea and along lower margin of clypeus. 

Eyes: In two rows, from above anterior row 
slightly recurved, posterior row straight. AME 

l~:gs. 1-4. KeUira jparsomacuiaia gen. ct ip nov. Hololypc myW | fej& hfyj n#'U palp, dofSflh 2-4 Uf( palp. 2 
vcnnal; 3 prolatcral; 4 rcuolateial. Sccde lines = 0.5mm. 





Hgs. 5-K). Ketlira sparsomaculata gen. el sp. nov. Allotype female, 5 8: 5 body, dorsal; 6 right leg I, prolateral; 7 
larsal claw; 8 ehehcerae, maxillae and labium. Paratype female (SAMA N 19878), 9-10: 9 epigyne; 10 vulva, veniral 
Scale lines: Figs 5 - 6, 8 * 0.5mm; figs 7, 9 - 10 = 0.1mm, 


■ J 

diameter 0.20 AME : ALfc : FMF ; PLh » 1: 1: 
0.9 : Qfl. I mef spaces. AMfi - AME 0.8 ; AME - 
ALE 0.5 i PME - PME 1.55; PME - PLE 1.55 ; 
AME - PME I. I ; ALE - PLE 1. MOQ: amef iot 

width . posterior width ; length = 2.75 . 3.4 . 2.65. 

Chelicerae; Length 1.20, width 1.69, geniculate, 
almost glabrous, setae long (ca 0.6), shorter on 
retiomargins. Three rows of teeth on fang groove; 
three teeth on promargin, middle tooth largest, three 
teeth on reuomargin. middle and distal teeth about 
equal, median row of about eleven minute pointed 
teeth, placed posterior to middle looth of promargin 
row. Labium: length 0.30, width 0.70, emarginate 
posterio-ktterally, six short selae in recurved row 
across anterior hall' with one near anterior margin 
midway towards each lateral c\lge. Longer setae on 
anterior margin, few setae on posterior half. 
Maxillae: Length 0.75, width 0.63. slightly 
Converging, setae long. Sternum: Length 1.25, width 
1.22, slightly convex, setae long. 

Legs: (Table 1). Scopuia sparse. Spines: Leg I, fe 
d2 p2 r3 (2 on right fe), ti rl vH, mt v4; Leg II, fe 
d2 p3 rl, ti v8, mr v4; Leg HI, fe d2 p3. ti v4, mt 
p2 v4; I eg IV, fe d2 pi, li pi v3. mr p2 v4; Palp, 
fe d2 pi all distal, ti p2. Claws: Teeth small wj|h 
nfcout 6 on proclaw. 4-5 on rctroclaw. 

Abdomen: Setae of short to moderate length. 
Spinnerets: ALS basal segment OJ0. apical segment 
0.08, PLS 0.20, O.fW. 

Palps: (Figs 2-4) Tibia retrolatcraily with 
apophysis carrying a short triangular blunt-lipped 
spur. Embolus thick, short, slraighl for the most 
part along prolateral side of cymbium, then curved 
inwards with a short blunt tip. 

Oesrripiitwt ofaUorype (Figs 5—10): As in linlotype 
but for the following: Carapace length 3.51, width 
3.50. Abdomen length 5.10, width 3.80. 
Colour in alcohol: Abdomen ereaniish with larger 
black spots, 

Eyes: AME diameter 0.24. AME : ALE : PME : 
PLE = I : 1.04 : 0.92 ; 0.9Z Interspaces: AME - 
AME 0,67; AME - ALE 0.5; PME - PME 1.3; 
PME -PLE 1.5; AME- PME I; ALE -PLE 92 
MOQ: A met tor width: posterioi width: length ■= 
IH : 3.2 : 2.6. 

Chelicerae; Length 1.60, width 2.10, Labium: 
Ungih 0.36, width 0.80. Maxillae: Length 0,94, 
width 0.71- Sternum: Length 1.80, width 1.80. 

Legs: (Table I). Spines: Leg I, fe (3 on right), ti. 
r2; Leg H, fe (2 on right); leg IV, fe p2, mi p3 rl. 

Spinnerets: ALS basal segment 0-36. apical 
segment 0.08. PLS 0.30, 0.10. 

Genitalia: bpigynum a low mound, highest 
posteriorly, small fossa with weakly sclcroiiscd 
hood-like anterior margin (Fig. *)>. Vulva paired, 
copulatory openings inside hood, spermalheca 

leading to a curved receptaculuni shortly joined to 
a smaller reeeptaculurn (Fig. 10), 
Etymology; The specific name refers to the widely 
scattered spots on the abdomen. 

KeiHca sokoii K0L «ov. 

Hotofvpe: AM KS19I41, a. Holey plains Siair 
Park. 3K° 13' Ss L46° 51* E. Vict., 29. ill. |980j A. 

De*eription of Nolotype: Similar td K. 
spunofHuciitaia except as follows: Carapace length 
3.44, width 3.22. Abdomen length 3.90, width 120. 

Colour in alcohol: With blackish markings 
laterally on carapace. Abdomen with numerous 

Carapace: Raised but flattish above, highest in 
foveal region, fovea distinct. 

Eyes: Both rows slight tv recurved, AME diameter 
0.20. AME : AL F. : PME : PLE = 1 : 0.9 ; 0.S : 
0.9. Interspaces: AME -AME I ; AME -ALE 0.9 
; PME -PME 1.7; PME -PLE I fcj AME -PME 
0.9 ; ALE - PLE Q.S. MOQ: Anterior width ; 
posterior width : length » 3-0 : 3.4 : 2.3. 

Chelicerae: Length J .44, width 1-72. Rd remargin 
with four teeth, irregular median raws o! 8 - 13 
minute teeth at ranged in triangular area with apex 
pointing distally. Labium; l-engihO.36, width 0.76. 
Short setae on posierior half. Sternum: I ength 1,76, 
width 1.70. 

1 cgs; (Table 2). Spines: Leg 1 fe dl p2 r3. ti r2 
v8, mt pi r2 v4; Leg 1L fe dl p2 r2, ti rl v6, mi 
p2 r2 v4; Leg III, fe d2 p2, ti p2 r2 v4, nit oO r2 
v4; Leg IV, fe dl pi u v4, mt pi r3 <2 on right) v4; 
Palp, fe d2 pi, all distal, ti dl p2 rl, 

Palp: legal u in positioned more hastily in 
cymbium than in k sparxamuculata and enlarged 
retrotetetally at proximal end. Embolus extending 
further in prolateral side, distal half straighter. 
Tibial spuT longer and more upright on larger 

ttyniotogy: The species is named after the collector, 
A. SokoL 


Known only from the holotype, K. sokofi U 
separated from K, spat soman* la (a by the tlflttcr 
carapace, four retrolauval teeth on the fang margin, 
several ii regular lows of median teeth and abdomen 
with more numerous spots. Its habits arc no; 


The completion of this paper was supported by 
an Australian Biological Resources Study grant. 
Special thanks go to Dr D. C Lcc for his helpful 
comments on manuscript preparation. Alsu to Dr 
M. Gray and C. Horseman lAM) for rhe loan ot 
material and assistance. 






Figs 11-15. Keiliru xokoli gen. et sp. nov. Holotype male: It Abdomen, dorsal; 12 cheliccral teeth, left retrolateral; 
13-15 left palp; 13 ventral; 14 proJateral; 15 retrolateral. Scale lines: Figs 11, 13 - 15 = 0.5mm; Fig. 12 = 0.1mm, 



Table 1. Leg measurements o/Keilira sparsomaculata gen. et sp. nov. values for holotype male with allotype female 
in parentheses. 













2.85 (3.32) 
3.21 (3.68) 
2.32 (2.56) 
2.45 (2.84) 
1.00 (1.00) 

1.55 (1.84) 
1.69 (2.00) 
1.12 (1.38) 
1.15 (1.36) 
0.52 (0.74) 

2.58 (2.82) 
2.92 (3.16) 
1.70 (1.96) 
2.00 (2.24) 
0.49 (0.80) 

2.50 (2.92) 
2.92 (3.16) 
1.69 (1.81) 
2.15 (2.40) 

0.90 (1.12) 
0.98 (1.24) 
0.80 (0.95) 
0.88 (1.00) 
1.50 (1.52) 

10.38 (12.02) 
11.72 (13.24) 
7.63 ( 8.86) 
8.63 ( 9.84) 
3.51 ( 4.06) 

Table 2. 

Leg measurements 

of Keilira sokoli 

gen. et sp. nov. i 

of holotype male. 

























Hogg, H. R. (1903) On the Australasian spiders of the 

subfamily Sparassinae. Proc. ZooL Soc Lond. 1902 (2), 

(1914) Spiders from Montebello Islands. Proc. 

ZooL Soc. Lond. 1914, 69-92. 
Jarvi, T. H. (1912-1914) Das Vaginalsystem der 

Sparassiden. Ann. Acad. Sci. Fenn. (A) 4, 1-235. 
Kaston, B. J. (1980) "How to know the spiders". 3rd Edtn 

<W. C. Brown & Co., Dubuque, Iowa). 
Keyserling, E. (1880) Die Spinnen aus Amerika. 

Laterigradae. Nurnberg, 1880, 1, 1-283. 

Koch. L. & Keyserlinc, E. (1871-1883) Die Arachniden 
Australiens. Vols 1 & 2. (Bauer & Raspc Nurnberg). 

Latreille, P. A. (1804) Tableau methodique des Insects. 
Nouv. Diet. Hist, Nat. 24, 129-200. 

Main, B. Y. (1976) "Spiders"- (Collins, Sydney). 

Simon. E. (1880) Revision de la famille des Sparassidae 
(Arachnides). Act. Soc. Linn. Bord. 34, 223-351. 

Walckfnaer, C. A. (1837) "HiStoire Naturelle des 
Insects". Apteres. I. (Paris). 




by Michael J. Tyler 


A large collection of frog ilia from the Tertiary site of Riversleigh Station in northwestern 
Queensland, includes 190 specimens of Lechriodus intergerivus sp. nov. The new species is 
described, compared with the four extant congeners, and shown to be the smallest member of the 
genus. A revised definition of the ilium of Lechriodus is provided, and the significance of the find 
as a link between the geographically isolated extant species is discussed. The rainforest habitat 
preferences of all extant species suggest a similar environment at Riversleigh Station in the Late 
Oligocene to Mid-Miocene. 
KEY WORDS: Lechriodus, ilia, Leptodactylidae, Tertiary, Queensland. 




by Michael I. Tyler* 


Tyi £R. M J. U9H9) A new species ol Levhrtodus (Anura; l^eptodactylidae) from the Ternary ot Queensland, 
with a redefinition oi' the iliaJ characteristics of the genu*. Trvris, /? Sot S. Ausr, 11.3, 15-;t, 31 May-, iy*9- 
A large collection of frog ilia from the Tertiary site of Rivcrsleigh Station in northwestern Queensland, 
includes 190 specimens of Lechriodus tntergerivus sp nov, The new specie^ is described, compared with 
the font t-Mani uongenern, and shown to he the smallest member of the genus. A revised definition of the 
ilium of Lechriodus is provided, and the significance of the find as a link between the geographically isolated 
exunt species is discussed. The rainforest habitat preferences of all extant species suggest a similar environment 
at Rivcrsleigh Station in the Late Ohgocene BO Mid-Miocene. 

Ki \ ■ Wokns: Lechriodus, ilia, l-eptodsctylidae, Tertiary, Queensland. 


Lechriodus Boulengcr comprises a group of 
ground-dwelling leptodactylid frogs confined to the 
Australian Geographical Region. )ti addition to the 
morphological attribute"* that support its 
recognition, it is unique <n the nature of its 
distribution! of the five leptodactylid genera that 
have been reported to occur both in Australia and 
New Guinea, it is the only one which is represented 
hv mora species in New Guinea than in Australia 
(Zweifel 1972), 

McDonald & Millet (1982) clarified the 
geographic distribution of lechriodus m Australia, 
demonstrated that a previous record from north 
Queensland was in error, and confirmed that there 
is a major disjunction between the Australian and 
New Guinea populations. Thus, whereas other 
shared genera occupy the Cape York Peninsula of 
north Queensland (Tyler 1972), Lechriodus does 
not*, and in Australia it is confined to the eastern 
fceaboard along the Great Dividing Range between 
Ourimbah, N.S.W, and Brisbane, Qld. All that can 
be interpreted about such allopatry is that at some 
time in the past Lechhodus must have occurred In 
the intermediate area. 

Here I report a uew species of Lechriodus from 
rhe Tertiary. The material comes from Rivcrsleigh 
Station in northwest Queensland, which is 
intermediate between the distribution o\ the extant 
species. The finding is .significant for several 
reasons: it constitutes the link in the distribution 
of the extant species; it is the first record of fossil 
Lechriodus, and it is the first occasion on which 
an Australian frog fossil has been reported from a 
site that is beyond the modern geographic range of 
its genus. 

Department of Zoology, I iniversirv of Adelaide* C.P.O. 
Box 498. Adelaide. S.Aust. 5001. 

Materia) and Method* 

The material is deposited in museums and other 
collections abbreviated in the text as follows: 
Australian Museum, Sydney - AM; Department of 
Zoology, University ol Adelaide - AUZ; 
Queensland Museum, Brisbane - QM: South 
Australian Museum, Adelaide - SaM, and 
American Museum of Natural History, New Y.-i k 
— AMNH. Letters following the abbreviations arc 
departmental identifications. 

Comparative studies were based on the 
osteologies! collections of the Department of 
Zoology, University ol Adelaide, supplemented by 
skeletal material of New Guinea taxa borrowed 
from the American Museum of Natural History. 

Osteological nomenclature follows lyicr (1976). 
The following measurements were taken with dial 
callipers: ilial length - measured from the superior 
extremity of the dorsal acetabular expansion to (he 
distal end of the shaft; dorsal acetabular expansion 
to ventral acetabular expansion (DAh-VA b ) 
measured as the distance between their extremities, 
and acetabular fossa diameter measured at the 
proximal, external edge of the peripheral acetabular 

Fig, I, Oricmaiion ol ilium to hon/ouud plane durim; 
preparation of description*. 

AiJ measurements and descriptive features ate 
derived from rhe lateral surface. Expressions ol 
relative extent of features arc obtained with the 
ilium positioned in such a way that the proxuiuil 
edge of the bone is maintained at 45 a to the 
horizontal (Fig. 1) Concepts of relative size of 



component features to one another are perceived 
with regard to their proportional size in other 
anuran taxa. 

Scanning electron micrographs were taken on an 
ETEC Autoscan SEM at 20 Kv. 

System atics 

Family: Leptodactylidae Werner, 1896. 

Sub-family: Limnodynastinae Lynch, 1971 
Genus: Lechriodus Boulenger, 1882 

The definition of the generic characteristics of 
the ilium by Tyler (1976) was based solely on L. 
melanopyga (Doria). Now that representatives of 
all extant species and the fossil species have become 
available, the definition requires modification and 

Ilial shaft slightly to moderately curved, bearing 
large, fanlike dorsal crest extending along at least 
three-quarters its length. Maximum depth of dorsal 
crest approximately one-third from proximal end. 
Crest concave to varying extents proximally, 
flattened distally. 

Acetabular fossa diameter varies with size of 
individual from which it is derived: proportionally 
larger in larger species. Acetabular fossa bordered 
by rim in large species; rim confined to inferior half 
in smaller species. Location of dorsal margin of 
acetabulum ranges from base of ilial shaft to 
midway up shaft. 

Pre-acetabular zone narrow, meeting inferior 
border of ilial shaft in gentle curvature, commonly 
forming quadrant. Ventral acetabular expansion 
moderately developed, commonly truncate 

Dorsal prominence small, extending superiorly 
above superior margin of ilium, or not. Dorsal 
prominence moderately to well developed, oval and 
horizontal or vertical. 

Dorsal acetabular expansion acutely angled, its 
superior margin on a level with or above the 
maximum extent of the dorsal crest. 

Lechriodus intergerivus sp. nov. 

FIG. 2 

Holotype: QM F16614. An almost entire left ilium 

collected at C.S. Site, Riversleigh Station, 


Description of holotype: Ilial shaft slightly curved 
and bears enlarged, fanlike dorsal crest whose 
maximum depth is at position approximately one- 
third from proximal end of shaft. Proximal one- 
half of crest concave on lateral surface, progressively 
becoming less pronounced distally. Distal half of 
crest flat. 

Acetabular fossa small, deep and with prominent 
rim bounding inferior half. Dorsal margin of 
acetabular fossa situated slightly superior to inferior 
margin of ilial shaft. 

Fig. 2, 

S.E.M. of Lechriodus intergerivus (SAM P29771). A left ilium, but image reversed to aid comparison with 

ANtu vHLCtKsni ticitMoi)US(\UVkr\ LEPIOOAGTVLMMTi 


Prv-acetabular zom: evenly rounded with narrow 
separation from acetabular fossa. Ventral acetabular 
Union poorly developed and incomplete 

Dorsal acetabular expansion narrow but well 
developed superiorly, extending to position 
approximately on a levcf with maximum extent of 
superior margin ol Ilia] GrtSL 

Dorsal prominence pocvly developed and scarcely 
detectable; in profile visible as slight superior 
cvlcnsion upon dorsal mgrein ot ilium. Dorsal 
protuherauec ovoid, lateral, well developed and 
projecting laterally. Anterior margin of dorsal 
prominence extends to position slightly anterior to 
ante riot margin of acetabulum , 

Length of ilium 13.6 mm; DAE-VAE 3.7 mm; 
acetabular fossa maximum diameter 1.4 mm. 

Paratypes: HenVA Hollow Site: SAM P29742; Two 
Trees Site: AM F7693W2, AMNH 25351-53; Last 
Minute Site SAM P297fS4-65; Gag Site- AM 
F76957. AMNH ID?5-Sft SAM P27%*; Upper 
Sitft AM F76953-54, 7f.95.S, AMNH 25354, QM 
l'mn40-50. 17036-39, SAM P29734, 29743-44, 
29757-W, C S Site . AM F76955-56, 76959, 
AMNH 2535^-60, QM rt6615-18, 16674, 
17031-32, SAM P2V746-50; Wayne* Wok Site; AM 
{76960, QM FI6636-J9* 17034-35, 17040-47. SAM 
P2975& 29766-67; Outasite Site; SAM P27929. 
2'>75l-55; RSO Site. QM PI66I<^35, 17033, SAM 
P2973S-4I, 29745, 29771 (subject of S.E.M | 

Variation: The complete ilia range in length from 
um DO 14.4 mm, A paratype is shown in Fig. 
2, I litre is minimal difference in overall shape, and 
variation is largely confined to flic position of the 
dorsal prominence in relation to the acetabulum 
Thus some specimens agree with Ihe holotype in 
Having the Ulterior margin ol the prominence on 
a level with the anterior margin of tie acetabulum, 
hi others the anterioi half of the prominence may 
project beyond the acetabulum. 

The extent and degree ol the curvature of the 
doisal eresi vanes m teftOS ol Its elevation, proximal 
Ifmi Stud medial protrusion, 

Rqfefrvd specimens; SomU pot nuns of an additional 
79 uperfrrjtftf m ^ilftaeritly complete to permit 
Idenafe^rton, bUl i0 fragmentary thai ihcy do nor 
lbUt€ to an understanding of ihe nature or 
variability ol Ihe species. For those reasons they arc 
considered here 'Vlcued specimens" rather than 
paraiype*). All of the referred specimens arc lodged 
111 tins QlUXn$tand Museum. KCTlkl Hollow Site: 
FI6652; Two Trees Site FI66K8-S9; Lasi Minute 
SilC I L6G7J, 16675, 1667K-79. 16685; Gag Site 
I I6S5I. H 6676- 77. 16680-83, lr>6S6>-K7 ( JgTOD-Ol; 
Uppcj Site; FJ$658-59, t66W-72, L7QOT-53;CJS 
Site f l6b60-64. 16684, 16690. 17048; R.S.CX Sitr' 

Ft665*-57. 16691-99, 170*9; \Vay.u'\ Wok Sftf 
F16782-14. I7U54 : 

Comparison w/Wi other spews. Ilia ol all M41U 
species have been examined; L fleichcn t Roulcnf cr) 
(AUZ 8 uncai. ). L, melunopyga (Dana) lAMNIl 
H122X AUZ 2 uii-.aLi; / aganaftoxis 7wcifel 
(AMNH M645), /_, platyceps Parker (AMNH 
74178). Representatives art shown in f i,c. 5. 

The comparisons ,:;v,i. n i.; ihat I hi- \ample 5iz*Ql 
L auergcrtvus is Mtlfiuehi k. rcflco with reasonable 
confidence the size ol Qph attained by the Kpei 1 
With that assumption it is relevant to npre rhcit the 
absolute sizes of the Ilia examined oi ettfi &l I 
adults or New Gumcn species is ^.ubMantialry lii/ger 
than any of the £, intent/ ivus. 

The relationship between mal length and Mil 
to venf length ol extant species is almost linear. 
Thus given ilial length alone it is porn 
extrapolate snout ro vem length, Ijil lot^catllium 
of/.,, intergenvur, r- 14.4 u ,n, (tic corresponding 
snout to vent length derived from the closely lineal 
relationship ol congeners is approsirnaMy 3 P.3mto 

(Fig. 4). This, the apparent rmtfmuni is 

two-thirds of the size ol (he smallest aduh / 
aganoposw and L plonite\ and is shinty tttlow 
the range ot'/.. mefawiw :*m\ I /- 1 

1) Clearly 1. inters nvws is the smallest member 
9f 'he gentifc 

As indicated by ihe generic definition, tfie ilium 
m this geuux is conservative and inlcrepcrific 
variation is not pronounced. For ihat reason I am 
reluctant to ascribe differences in EllaJ character* 
greater significance than iliey may tneilt- Howei 
given the data on size, the integnty of / . tn(rry,i'nvu\ 
is nol in nuesiion. 

Stratigraphy and lithotomy: The assumed 
stiatigraphic sequence ot the sites at Rivertfcijh i& 
that followed in the list of paratypes. Ot the wins 
romed to containing Z, inre^ensu-; I 
an almost uninterrupted wrjUCWft 0ue6l'Khc fossil 
bearing sediments is desaibed by Ffond (1985) te 
rine-grained arenacenus fresliwaier irmucones, and 
possibly a facics witriih (he ( . 1 - l 

The ages of the sed.rnenK cr-ntauung L tnwr^i ■ 
are currently understood 10 be between UdC 
Oliuocene and Middle Miocene (M. AicIil-/ 

Etymology: Lathi lor 'placed between". SO .lihuhnw 
to the ycographic position .m* tin; fojiij populS i 
bclwccn those ol ihr #£twil k iucir,<. 
Phytogeny 7weifers (ll>72) proposc-J fiiylngenctic 
nclationships is reproduced here as t (g, s. t;,iv C nthe 
geographic locaiion oi c. miergerivhs fRg, 6] I 
a^eorthcRivecsleieJi Stall , tid Acftun 

lhat the tower evtirtuitioii ot a^e is cojiretn r , 
■wuh the emcrjgcjKe and Nubsdquimi Mplift oi Hi ■ I 



Fig. 3. Ilia of extant species with or without ischium and pubis: A. Lechhodus fletcheri (AUZ uncat.); B. L. melanopyga 
(AMNH 81223); c. L. platyceps (AMNH 74178); D. L. aganoposis (AMNH 74646). 



1Q 15 20 

llmm Length (mm) 

tig. 4. Length of itia of Lechriodus species plotted against 
snout to vent length. Assumed snout to vent length of 
fossil species indicated by broken linev t-value for slope 
10.891, p < .001. For x = 14, y = 37.51 (95W ( . 
confidence umiu = 33.75-4J.25). Square = L, 
platyceps, circles = L. metanopygoi triangles = /.. 


Fig. 5. Phylogenctic relationships of extant species of 
Lechriodus as postulated by Zwcife! (1972), 

of New Guinea, L intergerivus is a candidate as 
the ancestral stock from which the extant species 

With the existence of L. intergerivus, ZweifePs 
phenetic scheme would require two invasions into 
New Guinea: one deriving L. melanopyga from an 
intermediate ancestor that also gave rise to /,. 
fletcheri, and the other leading to L, piatyceps and 
L. agonoposis. A slightly simpler scenario would 
derive L. fletcheri directly from L. intergerivus, with 
a second stock giving rise to the three New Guinea 
species, Further systematic studies are required to 
test the validity of that option. 


The fossil frog fauna of Australia includes 22 
species and nine genera from 10 sites of Quaternary 
or Tertiary age (Tyler 1989). Riversleigh Station 
appears likely to be the richest fossil frog site yet 
discovered, in terms of the number and diversity 
of species. This trend is common to other vertebrate- 
classes, and Archer, Hand & Godthelp (1988) stale 
thai more than 200 new species have been recovered 
there. The frog fauna will increase the total. 

Amongst the frog material from RJversleigh 
Station now available, L. intergerivus is the 
predominant species. Of 379 ilia now known from 
the site 190 (50%) represent that species. 

Extant Lechriodus are predominantly inhabitants 
of temperate and sub-tropical rainforest and, in the 
absence of any data to the contrary, it can be 
inferred that the habitat of L. intergerivus would 
have been rainforest. A second point relevant to 
interpreting the mid-Miocene environment is the 
fact that a high proportion of the ilia of other frog 
species found in Riversleigh Station deposits are 
from small creatures. Because frogs lose water 
readily through the skin, the body mass/surface 
area ratio is such that small frogs are particularly 
vulnerable to dehydration. In consequence they are 
predominant only in areas which are moist 
throughout the year. In communities of frogs in 
Australia high frequency of small frogs occurs only 
in areas of high and seasonally reliable rainfall, such 
as \hc extreme southwest of Western Australia and 
the northern periphery of Arnhem I .and in the 
Northern Territory (Tyler 1989). 


This investigation was made possible as a result 
of a suggestion from, and the subsequent 
encouragement of. Dr Michael Archer. I am further 
indebted to him for provision of laboratory space, 
and various help from his colleagues Mr Henk 
Godthelp and Dr Suzanne Hand at the University 
of New South Wales. 

The research program was funded by the 
Australian Research Grants Scheme, Veronica Ward 




l & 





• L rnelano pyga 

O L ag anoposis 

■ L platyce ps 

a L. fletcheri 

Fig. 6. Geographic distribution of Lechriodus. 



TABLb 1. Size o/Lcchriodus species in millimetres. Snout to vent length data mostly obtained from Zwcifcl (1972). 
Snout to vent length of donor L. aganoposis unknown, and maximum length of L, intergerivus extrapolated from 
Fig. 4. 




largest i 

Hum examined 

size of 
























played a vital role in that she undertook the 
cataloguing and initial sorting of material, and 
prepared Figures 1-2 and 4-6. 

Dr R. G. 7.weifel and Or C. W. Myers (American 
Museum of Natural History) lent tint of extant 
species and provided data about them, and Mr J. 
I. Menzies (National Museum & Arl Gallery, 
Konedobu) provided distribution records from 
Papua New Guinea. Mr P. Kempster prepared 
Figure 3, 

I am also indebted to Dr Keith Walker and Dr 
Margaret Davies for valuable discussions, and to 
the University of Adelaide for the provision of 
research facilities. 

The materials upon which this study was based 
were obtained through the support of the following 
funding bodies and organisations to M. Archer, S. 
Hand and H. Godthelp; Australian Research Grants 
Committee; Department of Arts, Spon, the 
Environment, Tourism and Territories; National 
Estate Program Grant Scheme; Wang Computers 
Pty Ltd; ICI Australia Pty Ltd; Australian 
Geographic Society Inc.; Mount Isa Mines Pty Ltd; 
the Queensland Museum; the Australian Museum; 
the Royal Zoological Society of NSW T ; the Linnean 
Society of NSW; Ansett/Wridgways Pty Ltd; 
Mount Isa Shire Council; the Riversleigh Society 
and the Friends of Riversleiuh, 


Archer, M., Hand, S. & Godthelp, H- (1988) A new 
Order of Tertiary zalambdodonl marsupials. Science 
239, 1528-1531. 

Hand, S. J. (1985) New Miocene megadcrmatids 
(Chiroptera: Megadermatidae) from Australia with 
comments on megadermatid phylogenetics. Aust. 
Mamnt. 8, 43-54. 

McDonald. K. R. & Miller, I D, (1982) On the status 
of Lcchriodus fletcheri (Boulenger) (Anura: 
Lepiodaetylidae) in northeast Queensland. Trans. R. 
Soc, S, Aust. 106, 220. 

Tyler. M. J. (1972) An analysis of the tower vertehrate 
faunal relationships of Australia and New Guinea. In 

D. Walker (Ed.) "Bridge and barrier; the natural and 
cultural history of Torres Strait". Dept of Biogeography 
& Geomurphology, Research School of Pacific Studies* 
Australian National University, Canberra. 

(1976) Comparative osteology of the pelvic girdle 

of Australian frogs and description of a new fossil 
genus. Thins. R. Soc. S. Aust. 100, 3-14 

(1989) "Australian frogs", (Viking O'Neil, 


Zweifel, R. G. (1972) A review of the frog genus 
Lechriodas (I^eptodactylidac) of New Guinea and 
Australia. Am, Mus, Novit, (2507), 1-41. 




Eight potential refuges for macroinvertebrates were sampled in two intermittent streams in central 

Victoria, Australia, during summer 1982-83 and summer 1983-84. Ninety-one aquatic taxa, mostly 

insects, were recorded. Receding pools harboured nearly three-quarters of these taxa; comparatively 

few were collected from the hyporheos or the water in crayfish burrows. Almost half the taxa were 

from refuges that did not hold free water. Macroinvertebrates persisted as desiccation-tolerant eggs 

(mayflies), larvae (chironomids and some beetles) or adults (beetles). 

There was remarkable similarity between the broad taxonomic representation in these refuges and 

those described for intermittent streams in Ontario, Canada. 

KEY WORDS: Intermittent streams, over- summering refuges, aquatic macroinvertebrates, Victoria, 

Australia, pholeteros, hyporheos. 


by A. J. Boulton* 


Boulton, A. J. (1989) Over-summering refuges of aquatic macromvenebrates in two intermittent streams 
in Victoria, Trans. R. Soc. & Aust, UX 23-34, 31 May, 19S9. 

Eight potential refuges for macromvcrtebralcs were sampled in two intermittent streams in centra! Victoria, 
Australia, during summer 1982-83 and summer 1983-84, Ninety-one aquatic taxa, mosily insects, were 
recorded. Receding pools harboured nearly iluee-quarters of these ta\a; comparatively few were collected 
from the hyporheos or the water in crayfish burrows. Almost half the urn were from refuses that did not 
hold free water. Macroinvertebrates persisted as desiccation-tolerant eggs (mayflicsk larvae (chironomids 
and some beetles) or adults (beetles). 

There was remarkable similarity between the broad ta^onomk representation in these refuge* and thave 
described for intermittent streams in Ontario, Canada 

Key Words Imerminem streams over-summenae refuges, aquatic macromveiTebraies, 
Vieioria, Australia, pholeieros, hypotheos. 


Ephemeral (episodic) and intermittent rivers and 
streams drain over half of the Australian mainland 
(W,D, Williams 1983) but despite their ubiquity and 
scientific interest, these systems have attracted little 
limnological attention (Boulton <& Sutcr 1986; 
Boulton* Lake 1988), The situation fe little better 
elsewhere (Williams 1987). 

[n intermittent streams, loss of water during the 
dry season is probably the most influential 
environmental parameter affecting the aquatic biota 
and has led to a wide range of physiological and 
behavioural adaptations (reviewed by Williams 
19S7). Behavioural avoidance appears to play a 
major part in the survival of many stream 
invertebrates during drought. Williams A Hyncs 
(1977) recognized eight distinct types of refuges thai 
were used by the fauna of a temporary stream in 
Ontario during summer and suggested that 
members of certain major taxonomic groups tended 
to over-summer as similar stages in their life cycle. 
Tor example, Ephemeroptera and most 
Chironornidae ovcr-summcrcd as eggs wherea* 
Gastropoda, some Odonata, HemipteTa and 
Coleopiera survived the dry pctiod as adults 
(Williams & Hynes 1977). 

in temperate Australian intermittent streams, 
recolonization pathways and potential over- 
summering refuges have never been investigated and 
little is known about the physiological or 
behavioural adaptations exhibited by the aquatic 

* Centre for Stream Ecology, Department of Botany and 
Zoology, Monash University, Clayton, Vic. .MfiS. 

Present Adfttvss; Department or Zoology, University 
of Adelaide, G.P.O. Bot 49B. Adelaide, S. Ami. 5001 

biota (Boulton & Lake 1988), This study was aimed 
at elucidating over-summering strategies of aquatic 
macroinvcrtebrates in two intermittent streants in 
Victoria. 1 also was interested to sec how closely 
ihe strategies used by biota in the intermittent 
Victorian streams matched those described by 
Williams St Hynes (1977) based on work done on 
intermittent streams in the northern hemisphere. 

Material* and Methods 
StVtty Arw 

Two study sites were located on the upper reaches 
of the Werribee River and two more on its main 
tributary, the Lerderder& Rivet (Fig. 11. Both rivers 
arise on the southern edge of the Great Dividing 
Range approximately J00 km north-west of 
Melbourne and flow south-east before joining near 
Bacchus Marsh, north-west of Melbourne. Details 
bf How regime and catchment vegetation are given 
in Boulton & Smith (1985); other physicochcmical 
data are presented in Boulton & Suter (1986}. 

The two rivers differ in permanency; the Werribee 
River ceases flow almost annually whereas the 
Lcrdcrdcrg River flows throughout summer for one 
year in three. On average, the Werribee River doe* 
not flow for nine weeks while the Lerdcrderg ceases 
How for Six. The Wertibee River dtd not flow at 
all at one site (Spargo Creek , SC) during the 1982 
drought and only flowed for five months (late June 
to late November) at the site downstream (Werribee 
Picnic Spot, WPS), The study pool at WPS dried 
up completely during the ensuing summer. The 
following year, flow commenced in late June and 
continued for seven and a half months at bocli sites. 

How started in late May 1982. at both siles 
(Fireplace Ford, FF and Wheeler Road, WR) on the 



Lerdcrderg River and ceased in early December. At 
FF, the study pool dried to a moist patch of leaves 
and water did not appear in the depression until 
mid-March 1983. At WR, the study pool was dry 
for six weeks before it also refilled in mid-March; 
flow resumed at both sites in early May, continuing 
over the following summer. 

Physicochemieal Monitoring of Potential Refuges 
On 2011983, diel ranges of temperature in and 
around a receding pool at WPS were measured 
using a 9-channel Miniature Intermittent. Temper- 
ature Recorder (Grant Instruments, Cambridge, 
England). Thermistors were placed in shallow (5 
cm) and deep (45 cm) water, below leaf litter^ 
beneath a large Hat rock, in the water of a crayfish 
{Engaeus sp.) burrow and in exposed grass in direct 
sunlight (regarded as "air temperature^ cf. normal 
meteorological practise). Recording commenced at 
5.00 a.m. and ceased at midnight. 

At other times, spot water temperatures (mercury 
thermometer), dissolved oxygen (oxygen probe. 
Model 51 A, Y.S.I. t Yellow Springs, Ohio), 

conductivity (conductivity meter. Radiometer, 
Denmark) and pH (Metrohm pH meter, Model 
CH9I00, England) were measured when potential 
refuges containing free water were sampled. 
Conductivity data were converted to values at 18°C 
(K 18 ) (Bayly & Williams 1973) whereas dissolved 
oxygen was expressed as percentage saturation using 
the conversion table in Bayly & Williams (1973) and 
an appropriate correclion factor for altitude. 

Biological Sampling of Potential Refuges 

A variety of collecting techniques was necessary 
to sample the diverse range of potential refuges: 
(a) An F.B.A. pond-net (300 /an mesh) was used 
to sample fauna in the receding pools. I vigorously 
shuffled along the bottom of the pool, sweeping 
the net from side to side across the disturbed path 
for 30 seconds for each sample. The size of the pool 
limited the number of samples that could be 
collected; while I was keen to ascertain the relative 
abundance of the fauna, 1 did not want to deplete 
the remnant populations. A nearby permanent lake 
(Shaws Lake, Fig. 1) was sampled similarly. 

To Trentham 



2^27 u 

Fig, 1 Map of the. study area showing I he locations of the tout studv sites: Wernbee Picnic Spot (WPS) - 1 
Spargo Creek (SC) = 2, Fireplace I'oid (FF) = j, Wheeler Roatl (WR) = 4. 



was recorded This proems was repealed until no 
fulDftt specks were seen. Subsequently, samples 
were taken ai l| regular intervals over the next 
fortnight, always reiurning the specimens to the 
tank. Some aquaria were maintained tor several 
months to rear hatch lings through to adults to assist 

were identified as far as practicable (sec 
Acknowledgments), Abundance was expressed 
qualitatively as "present*' (1-2 individuals), 
"common" (3-10) or "abundant" (>10); given the 
variety of collecting methods and the uneven 
sampling effort, more precise quantification was 

Physicwhemiwt conditions in potential refuges 

Means and ranges of spot water temperature, pH, 
dissolved o\ygen and conductivity in iwo refuges 
that held free water when they were sampled are 
listed in Table 1. Not surprisingly, the range*- of 
these variables were greater in the receding pools 
than in the burrow water of crayfish (Table 1), 
Hyporheic water was loo disturbed during sampling 
to obtain reliable physicochemical data and data 
from Shaws Lake are too few to be useful. 

Continuous records of wafer temperature In a 
receding pool at WPS illustrated the diel fluctuation 
or temperatures in vatious refuges (Fig, 2), Air 
temperature in direct sunlight near the pool ranged 
from 3.5°C ai dawn to 35°C early in the afternoon 
(Fig. 2>. The day was fine and clear with a light 
south-easterly breeze starting at 2.30 p.m. Sunrise 

Tari r I. Means and range* of water temperature, />//, dissolved tav^en and eomJuc/h'iry in two potential 
oven urn means refuyes, based upon n spot measurements 

(b) Animals residing in the water in crayfish 
{Engacus sp.) burrows ("pholeteros" sensu Lake 
]V77) were collected by carefully excavating the 
burrow opening and lowering a flexible plastic tube 
(6 mm internal diameter) into the burrow water 
which was sucked out into a plastic bag. In the 
laboratory, this water was sieved (50 /tm) with 
frequent washing to remove fine silt, 

(c) The hyporheo* was sampled by digging holes 
in the dry stream bed and sweeping a small hand- 
net (50 nm mesh) through the seepage. In some 
eases, it was necessary to use a plastic tube to collect 
(be water. Quantitative sampling of this habitat was 
not attempted. 

(d) Other potential refuges that were qualitatively 
sampled while the streams were dry included the 
humid mierohabitats beneath rocks, stumps and 
mats of leaf litter and dried filamentous aigae, and 
among the roots of riparian plants (e.g. Carex spp_, 
Leptospermum lanigerum) and stranded 
Myriophyllum propinquum (water milfoil) and 
Trig/ovhin procera (water ribbon), Rotting wood 
was broken open and examined, and strips of bark 
were peeled from exposed water-logged snags. 
Pieces of decomposed wood were brought back CO 
the laboratory for microscopic inspection 

(e) Plastic bags were filled with surface (upper 
10 cm) substrata and organic matter collected from 
the dry beds of riffles and pools. Tbe water eontenl 
of a subsample of the substratum was determined 
by subtracting the oven-dry weight (constant weight 
achieved after 48 h at I00°C) from the initial weight 
and expressing the value as a percentage. It was 



Tempc rat ui c 




f% saturation) 














4 7-7 2 














assumed that all weight lost during incubation was 
due to the evaporation of water. 

The rest of the sample was emptied into an 
aquarium immediately upon return to the 
laboratory and Hooded with dechiorinated tap 
waier, Within 30 minutes uf immersion of the 
substrata, a hand-net (50/im mesh) was swept 
vigorously through the tank and the live contents 
examined using an Olympus s>rereomicroscope, 
and the rank abundance of all inter icbraies present 

was at 6.21 a.m. but the pool was shaded by 
surrounding forest until about 9.00 a.m. Sunset was 
at 8.41 p.m. 

Water temperature in the shallows (5 cm) lagged 
closely behind air temperature (Fig. 2a) while in 
deeper water (45 cm) the daily range was far Ie*>5 
(Fig. 2b). Water in a crayfish burrow exhibited st 
dtel range of less than FC (Fig, 2b). The insulative 
capacity of several centimetres of eucalypt Utter and 
dried filamentous algal mat approximated that orf 




Buipjw wfoJfft 






2400 0600 



Ki£, 2 Variation m water temperature of several potential over-summering refuses in and around the study pool 
at WPS on 20.1.1983. See text tor details. 

a fiat rock 15 cm thick (of. Figs 2a and b). During 
the day, conductivity remained at 270 ^S/cm and 
pH ranged from 5.5 to 6.1 (n = 6 determinations, 
mean pH = 5,6). 

Aquatic macroinvertebrates collected from potential 

A total of 91 taxa was collected torn the eight 
over-summering refuges sampled at four sites on 
the Werribee and Lerderderg Rivers (Table 2). 
Nearly three-quarters of these taxa over-summered 
in receding pools along the river beds. These pools 
also harboured tadpoles of Litoria ewirtgi (Dume'ril 
& Bibron) (Hylidae) and Neobairachus sudellt 
(lamb) (Leptodactylidae); large numbers of the 
latter perished when the pool at WPS dried up over 
the summer 1982-83. Several specimens of Guluxiax 
olidus Gunthcr (Pisces: Galaxiidae) wore collected 
from the same pooL 

Few taxa were recorded from the hyporhcos or 
crayfish burrow water (Iable2); individuals in these 
habitats were usually tiny. Decomposed wood 
yielded small oligochaetes, tipulids and boring 
chironomids (Orthocladiinae) while amphipods 
(Austrochiltonia australis) were found in curled up 
strips of Eucalyptus spp. bark near tlte margins of 
dried out pools. These habitats were not sampled 
intensively and it is likely that other taxa (e.g. 
hclminthid beetles) may also use rotting wood as 
a refuge over summer. 

Almost half of the taxa recorded were collected 
from refuges that did not hold free water when 
sampled (Table 2). Some of these <e.g. Nousia spp. 
anil 'Winoioperta thmtitest) apparently hatched 
from desiccation-resistant eggs whereas others 

survived as larvae (chironomids, psephenids. 
hclminthids and case-building caddisflies) or as 
adults (dyliseids and helminthids) in moist 
microhabitats below rocks or mats of leaf litter and 
dried filamentous algae. Large numbers of 
microcrustacea (ostracods, harpaeticoid and 
cyclopoid copepods) were recovered from dry 
substrata flooded in the laboratory. The water 
content of these samples was usually less than 10% 
by weight (range = 0.19-74.00%, mean = 7.71%, 
S& = 0.93%, n = 53) and did not differ 
significantly between samples collected from the 
beds of the pool and riffle habitats (Mano-Whitney 
U-test, p > 0.05). 

Comparisons among the refuges should be made 
cautiously because the collecting techniques and the 
numbers of samples taken from each habitat 
differed substantially. For this reason, ] did not 
attempt calculations of coefficients of similarity of 
faunal composition among refuges, 


The classification of over-summering refuges 
used by aquatic macroinvertcbrates in temporary 
streams in Ontario (Fig. 3 iu Williams & Hynes 
1977) may be applied usefully to the data from the 
Werribee and lerderderg Rivers. At a broad 
taxonomic level, the faunal elements ovcr- 
summcring in the various refuges are almost 
identical and do so at similar stages of their life 
cycle. This may seem unexpected given the 
differences in latitude, source of stream flow 
(snowmelt in Ontario, rainfall in this study) and the 
difference in the degree of species overlap with 
nearby permanent streams (Boulton & Suter 19K6; 



Tam.e 2. Aquatic taxa recorded from potential refuges sampled over the 1982-1983 summer when the streams had 
ceased flow. Dead specimens were omitted. Abundance categories are qualitative (+ = present fl-)2 individuals/, 
C = common (3~\Q individuals), A = abundant {more than 10 individuals)) because of the different sampling frequency 
(n) and methodology (see text). * These individuals did not grow to identifiable size in the laboratory. ** (N\4V 
sp.n) refers to the number of the specimen held in the voucher collection of the Museums of Victoria. 



Below Crayfish Receding Shaws Wood Dry substrata Hyporheos 
dried {Engaeus sp.) pools Lake and flooded in 
litter burrows bark laboratory 

riffle pool 
(7) (12) (23) (2) (10) (47) (14) (5) 


IMesostoma sp. 

Cura pinguis (Weiss) 


Nematoda spp. 





Angrobia sp. 

Glacidorbis hedieyi Iredale 

Ferhssia spp. 

Physastra gibbosa (Gould) 


Sphaerium tasmanicum 



Oligochaeta spp. 



Heterias sp. 

Austrochittonia aus trails 


Paratya australiensis Kemp 

Cherax destructor Clark 

Engaeus sp. 





Limnesia spp. 

Austral Hobates spp. 

Corticacarus spp. 

Aquatic sp. 


Nousia spp. 
Atalophlebia sp. 



Austrolestes lio (Selys) 

Hemicordulia Itau Setys 







Crayfish Receding Shaws Wood 
{Engaeus sp.) pools Lake and 
burrows bark 



(2) (10) 

Dry substrata Hyporheos 

flooded in 

riffle pool 
(47) (14) (5) 



Acruroperla atra (Samal) 

Austrocerca tasmanica (Tillyard) 

IDinotoperla th wait est *Kimmins 



Microvelia dubia Hale 

M. distincta Matipatil 

Anisops deanei Brooks 

A. lhackeri Brooks 

Micronecta annae Uliesi 


M. a. tastnartica Wroblewski 


Antiporus blakei (Clark) 
A, femoralis (Boheman) 
Chostonectes johnsoni (Clark) 
Chostonectes spp. larvae 
Copelatus australiae Clark 
Hyderodes schuckardi Hope 
Lancetes lancealatus (Clark) 
Liodessus shuckhardi (Clark) 
Necterosoma penicitlatum (Clark) 
Necterosoma sp. larvae 
Piatynectes decempiwctatus 

Rhantus suturalis (Macleay) 
Sternopriscus mundanus Watts 


Macrogyrus sp. 

Hydraena luridipennh Macleay 
H, Itricantha Zwick 

Helodidae sp. larvae 

Sclerocypfion striaius 
Lea larvae 

Austrolimnlus hebrus Hinton 
A. maro Hinton 
A. "morrno" larvae (NMV sp. 
H 2 )** 

Austrolimnius sp, larvae 
Simsonia tasmanica (Blackburn) 

C C 











Limonia sp. 

Ormosia sp. 

Psychoda sp. (NMV sp. 3)** 






Receding Shaws Wood 

Dry substrata 



(Engaeus sp.) 

pools Lake and 

flooded in 




riffle pool 



(23) (2) (10) 

(47) (14) 



Aedes spp, A C 

Anopheles annulipes Walker C 

Culex fatigans Weidrnann C 

C. annulirosiris Skuse + 

C. austraiicus Skuse + 


Ablabesmyia sp. I (NMV sp. 

7E)** C 

Ablabesmyia sp. 2 (NMV sp. 


Paramehna spp. (nr NMV sp. 

32E)** C 

Chironomus nr februarius 

(NMV sp. 136E)** C + 

Einfeldia sp. + 

nr Dicrotendipes sp. (NMV sp. 

34E)** C 

Riethia sp. (NMV sp, SB}*' + 

Stenochirotwmus sp. (NMV sp. 

3E) + * 

Calopsectra sp. (NMV sp. 22E)'* C C 

StempeUina nr bausei + 

nr Monodiamesa sp. 

Orthocladius-Cricotopus complex 

(includes NMV spp. 12E and 

160E). C 

Heterotrissocfadius sp. \- 

Tiny chironomids* * C A 


Bezzia sp. 

Nilobezzia sp. 

Stratiomyidae spp. + C + 


Empididae spp. + C 


Dohchopodidae sp. 

Muscidae spp. C + 



Ptychobiosis nigrita (Banks) + 


Hellyethira Jsimpfex (Mosely) + 


Calocidae sp. 

Leptorussa dar/ingtoni (Banks) 

Oecetis sp. 

Led rides varians Mosely 

Triplectides similis Mosely 

T. truncatus Mosely 

Tiny leptocerids* 

Total number of taxa 91 
% of total number of taxa 























A. J. boi;i ION 

Boulton A Lake J9S8). However, physiological and 
behavioural adaptations employed by animals of 
eummon heritage thai share gross morphological 
similarities and that ate subjected 10 similar 
environ menial sciecttve pressures arc likely tu 
converge upon a restricted number of solutions 
(parallel evolution sensN Mayr 1963). 

In the Werxibee and Lcrderderg Rivers, several 
common laxa [Nousia sp., some gripopterygjd 
sloneflies) apparently hatched from deskcation- 
lcsistant eggs (cf. I^nmkuhl 1971; Snellen & Stewart 
1979; Malicky 1982) and were among, the fust 
iiwcrtcbtates to appear when flow resumed Similar 
findings have been reported in other ititeirauicnt 
streams (Harrison 1966; Chuiter 1968; Myites 1975; 
Laillc& Bass 1975; Williams & Hynes 19^6. 1977; 
Abcll 1984; Towns 1985k Although soTmhids 
(Dipteral were also common shortly afrer flow 
resumed in the Werribcc and Lcrderderg Rivera 
none emerged from the dry substrata Hooded in the 
laboratory, implying that eggs are Laid by adults 
flying in when flow .starts <cf. Hvnes 1975, Abell 

Mknicruslacea (ostracods. cyclopoid and 
harpacticoid copepods) emerged within hours of 
Hooding dry substrata in the laboiatory and 
harpacticoid copepods were observed mating a day 
later. Morton & Bayly (19771 recovered ovigenous 
harpaencoid females only 24 hours after flooding 
some dried mud from a temporal y pnnl ai Clayton, 
Victoria, suygesling that some species diapause at 
an advanced stage of development (cf. Cole 1953). 
Another taxon, common shortly after d7y substrata 
were inundated, was a neorhabdocccl Icntativcly 
identified as Mesostoma sp.. Bayly <I970) recorded 
Mesostorna from a temporary saline lake in south- 
eastern Australia and observed thick-shelled eggs 
in the uteri of some specimens lha'. are apparently 
teteased when Ihe .animals die, A similar strategy 
for desiccation-tolerance has been observed in 
ncorhabdocoets from a temporary ditch in Cnglnjid 
(Cm & Young 1974)- 

Gastropods in the Werribcc and Lerderdeig 
Rivers survive diOughi either by secreling a 
protective epiphruem (e.g. Ptiysastra gibbosa, 
FerrissM spp.)(cf. KcnJc 194$; Eckbtad 1973- Legier 
& Tab'n 1973) or by closing Their operculum (e,g. 
Angrolw sp. and Gloadorbis hedfeyf) (Boulton & 
Smith 1985). Musi lenJed to aestivate in moist 
miciohahiiats under stumps, dry algal mats and leaf 
liner (cf- Strnndine 1941; KlekowskI 1959; Casey & 
Ladle 1976). 1 he bivalve Sphaerium tastmanu-um 
probably minimizes water loss by closing its valves. 
Other members of the Spltaeriidae are 
ovoviviparous and brood their young while surface 
Onto is absent (Heard 1977; Hcinbacb etal 1980, 
McKcc & Mackie 1983). Activating juveniles have 

been found buried in the substratum (Way et ai 
1980) but I did not recover any from my siudy sites. 

PaTastacid crayfish found in the Werribee River 
over-summer in iheir burrows where water 
temperatures remain quite constant* Surprisingly 
few other imertcbrales (phnlcteros s*vuv Lake 1 97"*) 
appear to use this refuge (cf. Creasei 1931; Williams 
era!. 1974; Williams & Hynes 1976; Wiggins et at 
1980). The pholeteros in other Victorian ErigtHiuy 
spp* burrows is also depauperate (Horwity ev ai. 
1VH5). Neverlheless, this refuge seems to be 
important for the survival of janirid isnpods in the 
Werribee River. Isopods usually are absent from 
temporary waters because they lack desiccation 
resistant stages and are sedentary (Williams 1985). 
The water in burrows constructed by fish serves as 
sources of recoionists after droughts in some 
streams overseas (e.g. Tramer 1977; Glodck 1978) 
but this refuge was not evident in ihe Werribee and 
Lcrderderg Rivers. 

few laxa were abundant in the hypurheus of the 
study sites during summer. This paucity m&y reflect 
ihe crude sampling methods because amphipuilx, 
janirid isopods, stoneflies, molluscs and 
oligochaeies have been recorded from the hyporhcos 
of the intermittent Bxaehina River in South 
Australia (W.D. Williams 1983), Overseas, the 
hyporhcos is considered to be an important refuge 
from both droughts and floods in some temporary 
and permanent streams (Clifford 1966; Williams 
& Hynes 1977; Williams 1977, I9K4). However, in 
desert streams whose beds are mainly composed of 
unstable sand, hyporheic dormant stages are rare 
because of higj] tempeiatures in the dry strearobe d 
and severe scouring of the channel during flash 
floods (Gray 1981, Fisher et ai 1982). 

One refuge not considered by Williams & Hynes 
11977), possihly uncommon in Ontario, is that 
provided by decomposing wood debris, abundant 
along the banks and stream beds ol many Aust- 
ralian intermittent streams. In my study, this refuse 
harboured oligochaetes> araphipods, tipulids and 
chironornids. More intensive sampling is likely to 
vield further taxa other workers have 
recorded large numbers of xylophitous taxa From 
permanent streams in Oregon (Anderson of at 1978; 
Dudley & Anderson 19781 tinti New Zealand 
(Anderson 1982). 

In the Werxibee and l^rderderg Rivers, taxa 
aestivated under rocks and mats of algae (where 
temperatures remained constant) as adults (e.g. 
amphipods, dytiseids)or larvae (e.g. chironornids, 
stratiomyids). Some larval stages seemed 
surprisingly tolerant of desiccation; (or evampJe, a 
large water penny larva (Svterorypfjot\ strkrtush 
colle^ed from the exposed surface of a flat rocV 
on the bed of a riffle that had not flowed to? U 



weeks, resumed activity immediately after 
immersion m water from 4 nearby pool. Similar 
tolerance of desiccation by larvae of Scterocvphon 
spp. has been de&cribcd in Tasmania (Smith I9SI) 
and Queensland (Smith & Pearson 1985). 

Adult aquatic Hemiptera (e.g. cotixids, 
notoneetids* and Coteoptera (e.g. dyiiseidv> prob- 
ably fly tu from nearby permanent waters. Such 
aerial recxMonisatjon of temporary pools anil 
streams by these groups is commonplace {eg. 
Fernando 1958, 1959; Fernando & Gaibraith 1973; 
Williams AHynes 1976, 1977; Wiggins eiai, 1980. 
Abell 1984; Williams J9S5). I also found severaJ 
species of adult dytiscids over-summering, below 
rocks and dry Utter near the margins of pools at 
SC and WPS- This strategy seems less well-known 
for this group; only D.D. Williams (1983). 
Boumezzouah (1983) and McKaige (1980)' (in a 
temporary pond near Cotac western Victoria) have 
reported similar observations. 

Although the receding pools harboured most of 
the taxa that nver-surnmer at tbe study sites, they 
appeared to be the most physieuchemicalfy liansh" 
refuge that f sampled. The pools experienced a 
considerable d»ei range in water temperature, oxygen 
level* were frequently below 2OTo saturation and pH 
levels fell to less (ban 5. Conductivity rises as the 
water evaporates and the pools arc often stained 
dark brown wilh eucalypt leachate. Similar 
conditions have been observed in receding pools in 
other intermittent streams in Australia flbwns 1983, 
1985; Smith & Pearson 1987) and North America 
(Slack 1955; Larimore <?/ <rZ 1959; Clifford 1966; 
Harrcl k Don is 1968). 

The invertebrates that over-summer m these pools 
have various adaptations that allow them 10 tolerate 
sueh physicochemical extremes. For example. larval 
dyiiacids come up to the surface of the pool to 
obtain air through iJie terminal abdominal spiracle 
wheieas aduli dytisads store ait beneath their elytra 
(Britton 1970), Some chironomid larvae (e.g. 
CblronowtiS nr februarius) are particularly 
abundant in the receding pools and use 
haemoglobin to facilitate oxygen uptake (Collcss & 
McAlpbte 1970). Mayfly nymphs {Ataiophtebia sp.), 
also common In this refuge, have large sills that are 
constantly oscillated to enhance respiration 
(Uoulton & Lake 1388). Terrestrial oviposition by 
two species of* tsptoeerid caddisfly common in the 
Lerderderg River (Leptorussa duriingtofv' and 
Learxfes variant) may also be an adaptation to low 
or unpredictably fluctuating oxygen levels (Towns 
1983). Development of vulnerable juvenile stages 

' McKftige, M,E* (19.50) Emergence and development of 
*qtia1tc invertebrate communities from dried mud after 
flooding. ELSc Horn. rhefiis, Department of Zoologv, 
Motkish University (Unpubl.). 

lit a prrysicochemieally harsh environment may be 
avoided or accelerated by ovovivtparity exemplified 
by the stoneily Ausuvcerce iasmantea, also 
recorded by Towns (1985) in a South Australian 
Intermittent stream 

Although there U little known about the thermal 
tolerances of aquatic mactoinvertebtates in 
Australian intermittent streams, it appears dial 
many taxa can cope with short term exposure to 
extremes of water temperature It is likelv that they 
remain near the bottom of the pool where the 
temperatures may be as much as I5°C cooler than 
those of the surface water exernpbfied by the study 
pnnl at WPS. Such stratification is uncommon jn 
shallow pooh (eg. Byars I960; Butter 1963; Moore 
1970; Hartland-Rowe 1972). However, Erilcsen 
(1966) recorded surface-bottom differences of 
9-16°C in temporary turbid puddles less than 50cm 
deep, and a shallow (10 cut) rockpool in a stream 
in the Pyrenees had a surface temperature of 29.9°C 
while the bottom was I9°C fChodorowska & 
Chodorowski 1966), l^s.v marked stratification in 
pools in North American intermittent streams has 
been reported by Neel (1951) and Slack (1955). 

Isolated pools remaining in the stream bed are 
important over-summering refuges for aquatic 
macroinvenchrares in intermittent streams elsewhere 
(e.g. Slack 1955; Paloumpis 1958; La ri more et at 
1959; Williams & Hynes 1976; Abell 1984). In 
temporary streams in Ontario, they also provide 
excellent breeding environments due to the ease with 
which they warm up and the abundanr plain food 
that develops within them, and they enable species 
wilh long-lived aquatic stages to complete their fife- 
cyclcs (Williams & Hynes 1976). Similarly, in the 
Werribceand Lerderderg Rivers, these pools suppon 
a rich fauna although environmental condiions are 
harsh and predators are numerous (Boukoit & Suirr 

Tfuca whose aquatic life-spans are brief enough 
to he completed while water U present are under 
less selective pressure to adopt these strategies than 
species whose aquatic development takes longer 
than the period that water persists. Unfortunately, 
we lack information on the duration of aquatic 
stages of many Australian macroinvertebratcs and 
few generalisations may be drawn. Most crustaceans 
and molluscs listed Dl Table 2 probably live longer 
than a year and this may account for their ability 
to survive in refuges other than the ponfe. Although 
many other taxa (e.g. chironomids, culictds) can 
complete their aquatic stages in a matter r.f weeks, 
rheir survival in intermittent streams depends upon 
when their eggs hatch and how long water persists 
afterwards. Interpretation oi the significance Of 
many of these refuges will be possible when more 
information on the life histoiies of 



macroLnveitcbratcs in Australian intermittent 
si reams, is available. 

In summary, there appear \<y be five major over- 
summetlng strategies employed by the fauna of 
these two rivers; 

i) toleraring extreme and variable environmen- 
tal conditions in the remaining pools, 

ji) surviving in ?noi« nutrohabitais below stones, 
stumps and mats of dried algae and leaf litter, and 
in rotting wood, 

tii) over-summering in microhabiiats where 
environmental conditions are relatively mild and 
constant such as in the liyporheos or the burrow 
water of crayfish, 

iv) surviving as desiccation-resistant wages in the 
dry substratum, and, 

v) living in nearby permanent water-bodies and 
Hying in and ovipositing when flow resumes. 

Tne relative contributions 1'rom each of these 
refuges reflects the nature of the substnuum (e.g. 
Clifford \%6, Gray 1981), the amount and pattern 
of discharge during the previous spring and the 
seventy of the summer. This last was illustrated by 
the 1982 drought when all the pools in the upper 
reaches of the Werribee River dried completely, 
extinguishing seveial common tax&tog. the shrimp 
fttratya oustraUensh) that I never recorded the 
following year. Differential vurvival of fauna in 
these various refuges undoubtedly influences the 
community composition of the stream and has a 
profound effect upon its ecological succession 
during the ensuing period of flow. 


It Is a pleasure to acknowledge the enthusiastic 
encouragement aud guidance from Dr Sam Lake, 
Centre for Stream Ecology, Monash University 
where 1 did this research supported by » 
Commonwealth Postgraduate Research Awatd. 

Many taxonurnisu identified my 
macrolnvertebrates; I thank Dr M. Harvey 
(Hydtaearina). Or CMS. Watts (U>tiscidac), Ms 
T. Weir (Hdminthidae), DrJ, Davis (P<eph en idae)> 
Dr Ci. Theiscliinger and Ms C. Yule (Pkcoptcrai. 
Dr P. Suler (Ephemeiuplcra), Dt A. Wells 
(Hydroptilidac), Ms R. St Clair (Piatynelrninlhev 
Lepioceridae), Dr P. Iloiwiu (Pataslacidae), Drh 
Lansbury (Hemipter*), Dr* W. Fender and U. Smiih 
(Gastropoda) and Dr A. Sotool (Anura). Dr R. 
Marchanl, Mr L. Mctzeling anxi Mr, P. Lilywhltc 
allowed me access to laboratory facilities and the 
voucher collection at the National Museum of 

I was helped in the field by Iwn Barmuta, Mark- 
Harvey, Pierre Horwit?, Paul Humphries, Sabine 
Schrciber, Anthony SofcoK Gaye Spangaro, Alistair 
Suren and Cathy Yule and the publican at Spargo 
Creek deserves special mention for his provision oi 
an over-summering refuge for semi-aquatic 
vertebrates, Mr Dcvo Morton kindly supplied me 
with aquaria and other equipment, 

I am indebted to Dis Margaret Davies, Sam Lake 
and Keith Walker, Prof. Bill Williams and two 
referees for constructive comments on the 


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The larva of Odontacarus (Leogonius) adelaideae (Womersley, 1944) is redescribed, and a 
lectotype designated. Larvae have been collected repeatedly on domestic cats in suburban Adelaide, 
South Australia, and reared experimentally to protonymphs and deutonymphs. The larva, 
protonymph and deutonymph are described. Experimental transfer of larvae from cats to man is 
described. A key is given to the known deutonymphs of Leogonius, and revisional notes on the 
classification of the larvae. 

KEY WORDS: Taxonomy, Odontacarus, Leogonius, South Australia, larva, protonymph, 
deutonymph, rearing, cat. 




by R, V. Southcott* 


Sormtrorr, R, V (1989) The Ijirva and nymph msiarvof Odontacarus (Leogonius) adclatdeae ( Womersley) 
(Acarina: Irombieulidac: Leeuwenhoekiinae). Trans. R, Soc. S. Aust. 113(00), 35-45, 31 May, 1989 

The larva of Odontacarus (Leogomus) arfetaideu€ (Womersley, 1944) is rcdescribed, and a lectotype 
designated, Larvae IWVC been collected repeatedly on domestic cats in suburban Adelaide, South Australia, 
and reared experimentally to prolonymphs and deutonymphs. The larva, prolonymph and Jeutonytnpn 
are described. Experimental transfer ot larvae from cats to man is described. A key is given to the known 
deutonymphs of Leo^onms, and revisions) notes on the classification of the larvae, 

Key Words- Taxonomy, Odoatocara\ Leogonius, South Australia, larva, proionymph, 
deutonymph, rearing, cat, 


Womersley (1944) described Leeuwenhoekta 
adelaideae from several larval trombidioid mites, 
collected from domestic cats In southern suburbs 
of Adelaide in 1931 and 1941, and three collected 
from rats, in Cairns, Queensland, in 1939. He had 
writer (Womersley 1934) relet red to some of the 
same (1931) larvae as "Leuwenhoekia (sic) 
australiense (sic) Hlrsl 1929" (sic), i.e. as 
Leeuwenhoekta aust 'ralicnus Hirst, 1925, originally 
described from larvae collected at Ashfield. Sydney, 
New Soutli Wales, but subsequently recorded more 
widely in Australia and the south-west Pacific area 
(Cumber '952; Wharton & Fuller 1952), 

Following revisions of the taxonomy of the 
leeuwenhoekiine larval mites, (hose known from 
Australia, New Guinea and south-east Asia 
(previously referred to Leeuwenhoekta Oudemans, 
1911 and Acomatacarus Ewing, 1942) are referred 
to Odontacarus Ewing, 1929, subgenus Leogonius 
Vercammcn-Grandjcan, 1968 (see Southcott 1986a). 

In the present paper the larva of Odontacarus 
adelaideae is redescribed. The first attempt to rear 
larvae was in 1941, but the protonympbs and 
deutonymphs described below were not obtained 
until 1950-1951 

A key is given for deutonymphs of Leogonius* 
The successful transfers of pattly-fed larvae- from 
cats to feeding on man is described. 

Materials and Methods 

Slide-mounted mites in the South Australian 
Museum Adelaide (SAM) referred to O, adelaideae 
and collected by D. C. Swart in 1931 and R. V. 
Southcott in 1941 were examined. They were 
originally mounted ip gum-chloral media, hut 

Taylors Road. Mitchmn, S. Aust. 5062. 

showed evidence of extensive remounting to gum- 
chloral or polyvinyl alcohol-based media, with 
associated damage. Some specimens labelled as 
adelaideae from N.S.W, were excluded. The 
specimens' recorded by Womersley (1944) from 
Cairns, Qld, 1939 were not located, 

Further larvae were collected topotypically in 
1941 and between 1946-1952 from the ears of 
domestic cats. Some were mounted in water-soluble 
media (generally gum-chloral based), whilst 
attempts were made to rear others to later instars. 

In 1941 larvae freshly collected from cats were 
confined under a watch-glass attached to my 
forearm with adhesive strapping. Although the 
miles re-attached and increased in size, no successful 
transformation occurred. 

Mites were again collected from the ears of 
domestic cats from the same sire, during summer- 
auTumn of 1946-1952> and placed on strips of damp 
blotting paper tn sealed tubes at ambient 
temperatures. In one case a nearly-intact 
p-rotonymph was obtained. In several cww 
transformation 10 deutonymph occurred. All 
specimens were then preserved in lactic acid, or dry, 
until studied, generally 35-40 years later. The 
deutonymphs and residual pieces of larval or 
protonymphal pelts were slide-mounted, through 
lactic acid to Hovei's gum chloral medium (Baket 
& Wharton 1952*). 

Microscopy was by a LeiuOrttiolux microscope 
with phase-contrast and polarizing facilities; its 
camera lucida was used to make the line drawings. 

Terminology and abbreviations are as in 
Southcott (I986a,b). All measurements are in um 
unless otherwise stated. Prefixes: ACB to aurhor*s 
registration numbers, N to ihose of SAM. 

Odontacarus Hwing 
Odontacarus Ewing, 1929, p. 188 

(for synonymy see Southcott. 1986a. p. 171, aiiU 
contained references). 



R i 

"'"• -. '*^' I L III' 1 W Ss" 1 ^'-, 


vV'7 V^ 

CI "■ 





:a'3^i?r"r:---">^. '■:■■■■ ; 

Fig. 1 Odon iacat us acte/aideae (Womcrslcy), larva. A Dorsal view of leeLotype, legs on left omitted beyond uochanier*; 
Icgy on right completed from specimens in same postures as tectotype, leg f from specimen ACB520BA2, leg 11 
from ACB270A, leg III from ACH270B. B Gnathosoma, dorsal scutum and adjacent structures from paralectotvp^ 
N (988333 {AKtifyfy C Same for specimen ACR520BA2. D. Dorsal idiosomal seta. F Ventral idio^omal seta. I 
Tibia 1 and tarus I, dorsal aspect, of paralecrorype NI988333. (D, £ (o scale on right; oilier figures to scale on left I- 


Definition of (ana; Trombidioidca (as defined by 
Southcott 1987) with anterior median dorsal 
scutum, with riakllft, CWO anteromedian setae, two 
anteiolateral setae, two posterolateral setae, and two 
scnsilla* each with non-expanded sensillary seta. Leg 
tegmental formula 6, 6, 6, Eyes 2 + 2, sessile. 
Definition of deuionymph and adult. 
Trombidioidca with anterior median crista. 
expanded anteriorly to a blunt arrow-head-like 
-luipe (sa^illa), beat iffg two normal selae, with two 
-^aMlia towards ils posterior end, each bearing a 
slender* setulose sensillary seta, lxg segmental 
formula 7, 7, 7 Deutonymph with two pairs of 
suckers (aceiabula) along each lateral genital valve, 
adult with three pairs. Dorsal idiosomal setae not 
leal-like, nor divided, bui may be present in two 
dislinci forms, Eyes absent. 
Typespectex Trotnbicuh dentato Ewing, 1925, p, 

Subgenus Leogomus Vercammcn-Grandjcan. 1968 
Definition of larva: as m CtafC (1979, p. 143). 
TVpe species: Leeuwvnhaekia australiensis Hirst, 
1925. p. 150. 

Odonrucarus cdelatdeoe (Womersley) 
FIGS IA-E, 2-6 
lA'Uwunhoekia (sic) austrahense: (sic) Wbrncrslcy. 
1934, p. 217 fparnrn). 

Leeuwenhoekia autfwlieme; (sic) Womer$1ey T 1937, 
p. 82 (partim). 

U'euwenhoekia attsiraticnsis; Womersley &. Heaslip, 
J943, p. 141 (partim): Thor St Willmann, 1947, p. 
323 (parti m). 

Leeuwenhoekia adetaideue Womersley, 1944, p. 105; 
Gill, Moule & Riek } 1*45, p. 29; Gill & Parristh 
1945, Plate 3; Taylor. 1946, p. 128. 
AcomtitQcams adekiideae; Womersley, 1945, pp. 9S, 
III; Souihcotl. 1957. p. 149. 
Acomaiacarus ( Acomatacarus) adelaideae: 
Wharron & Fuller, 1952. p. 97. 
Qdonfanirus adeialdeax Southcotl, 1973, pp. 46, 
103; 1976, p, 139; 1978, p. 16 (unnamed); let &. 
Souihcotl, 1979, p. 35; 1980, p. 7. 
(Jdontovarwt (Leogomus} adefatdeae; Southcott, 
1986a. p 180. 

Dtrsxrtpiion of iarva: i^ectotype. Colour in life 
orange red. Length of idiosoma (mounted on .sltde) 
370. width 285; total length from tip of chehccral 
fangs to posterior pole uf idiosoma 463. 

Dorsal scutum wider than long (nasus included); 
nastrs well developed, tongue-like, slightly pointed 
anteriorly, slightly waisted, meeting body bf scutum 
at approximately right angles; anterolateral angles 
slightly obtuse, rounded, lateral borders slightly 
convex; posterolateral angles obtuse; rounded; 

posleiolcjlera] borders sinuou*; posterior angle 
obtuse, rounded. Sctitalac narrow, tapering, slightly 
blunted at tip, with pointed, auiMandlng sctules. 
Sensillaiy setae with 8-10 seiules in distal half. 
Sensillary sockets level with PL scutalae bases and 
&el slightly obliquely. Shield lightly poro^t, with two 
larger pits near each PL angle. 

Standard and other data of scutum and legs OX' 
the type scries as in Tabic J 

Eyes oval, conjoined, posterolateral to dorsal 
scutum; anterior eye with maximum diameter 16, 
posterior I J. 

Dotsal Idiosomalae normal, lightly tapering, 
slightly blunted at tip, moderately setulose, arranged 
2 rhumeraU"), lhen 6, then m vague rows across 
dorsum; loial about 55. 

Ventral surface of idiosoma wiiti a pair of 
pointed, setulose setae, 33 long, with centres of 
bases 34 apart, between coxae lit. Behind coxae 111 
opjsthosoma with 42 setae, curved,, setulose, 
pointed, but tbc more posterior setae longei, slightly 
btunted, resembling posterior dorsal idiosomalae 
Anus 24 long by J3across; 1 6 setae anterior to level 
ol middle of anus, and 26 posterior. Lrstigma well 
chitinized, oval, 25 long by 16 wide. 

Coxatae 2, I, 1. long, well setulose, tapering, 
pointed. Lateral coxala I 64 long, medial coxala 1 
68, ooxala II 53, coxala III 46, 

Dorsal tracheal opening present between 
posterolateral edge of gnathosoma and anterior 
border of coxa 1 (or overlying latter). 

Dotsal tracheal opening present between 
posterolateral edge of gnathosoma and anterior 
bonder of coxa 1 (or overlying latter). 

Gnathosoma normal. Combined chelicerae bases 
c, 7!? across, by 91 long from tip of cheliceral fangs 
to posterioi edge of cheliceral bases. Fangs stout, 
curved, blunt-pointed, with 3-5 strong retrorsc teeth 
along concave (flexor, dorsal) edge, ;tnd 5-7 blunled 
denticles along convex (extensor, ventral) edge. 
Galcala 26 long, simple. 

Palpi normal, palpal setal formula ft, fi, BNN, 
So + 711 Palpal tibial claw (brec-pronged. No 
supracoxala to gnathosoma or legs. 
Description of legs of larva (Horn ACB52UBA2 and 
ACB270A.8): Leg lenglhs (including coxae and 
claw*): I 425, II 355, III 425. Scobalar formulae: 
trochanters I, I, I; femora 6, 5. 4; genua 4, 4, 4; 
tibiae 8, 6, 6 (including 2 masialae on tibia 111); tarsi 
26, 16, 13 (including one mastala on MIL leg 
specialized setae as folluws: Leg I (from 
ACB520BA2* 5oGel.38ad(2l long). VstieL66pd(4), 
Sc>Gel.7|pd(22>, SoTiL59d<16), VVTil.88d(2)» 
Scil il.95adtlO) (Le slightly distal 10 Vs), Leg 11 (from 
ACB270A); SoGeJ1.34pd(20) J VsCeIL64pd(4), 
Solill.42pdU6;. SnTiH.S9pd(l0) Ug 111 (from 
ACB270B); SoGel!L26pd(2l), S6rilll.50pd(27). 





ft V"- 

v \);-|/ I 

^V> ;/ > 

Fig. 2 Odontacarus adelaideac (Womersley), larva. Ventral view of lectotype, legs on Icli 
oniiiTcd beyond trochanters; legs on right completed as in his.. 1. 

Table I. Metric data for type series of Odontacarus adelaideae {Womersley) larvae 




1 rectory pe 















+ 2.77 




























+ 2.83 















+ 1.64 





































Hum 1 















40.60 2 

























+ 2.83 












+ 3.46 












+ 2.83 





































(1.19- 1-24) 







1 Humeral seta length 

2 For maximum values 

Both tarsi I and II bear a large central dorsal 
solenoidala: SoTaI.43d(15) and FaTal.40ad(3) (from 
ACB270A); on tarsus II (from ACB270B) is 
SoTaII.40d(15), and FaTaII.37ad(4) (i.e. slightly 
proximal and anterior to Sofa 1 1 (in usual convention 
of legs being considered stretched out horizontally 
at right angles to main longitudinal axis of mite). 
Pretarsal formula I, 1, 0. Tarsal claws normal, 
slender, falciform, simple, neomedian longer than 
anterior and posterior, each with a fringe of minute 

Description of Proionymph: (Fig. 3) from mounted 
specimen ACB522R\5) Colour in life not observed, 
presumably orange-red. Shape irregularly 
spheroidal, 630 long, maximum width 430. Skin 
without .setae, thin, more or less smooth but with 
many minor creases, possibly artefacts of mounting. 

Developing limbs represented by finger-like 
processes; those of ehelicerae and palpi still adnate 
to main mass of protonymph, those of legs 
becoming free. 

A few setae and a few fragments of larval 
tracheae adhering to protonymphal skin. 

Description of Deutonymph: (Figs 4-6) (from slide- 
mounted specimen ACB522AA/DI, supplemented 
by other specimens) Colour in life orange, ldiosoma 
ovoid, more pointed anteriorly and posteriorly, not 
waisted, flattened below; no division beLween 
propodosoma and metapodosoma. Propodosoma 
more or less conical; posterior pole of idiosoma 
rounded, Length of idiosoma (from tip of sagitta) 
512, width 370; over-all length from ehelicerae tips 
to posterior pole of idiosoma 634. 



Fig. 3 Odontacants udelaideae (Womersley). Protonymph, 
entire, specimen ACB522BA5. 

Propodosoma bearing dorsalty a short crista 151 
long, with sagitta about 15 across, carrying I wo 
almost nude pointed setae 41 long, Sensiilary area 
46 across, wiih iwo large sensilta, each with a 
Filiform sensiilary seta 100 long, with faint setules 
in distal halt". Crista produced posteriorad beyond 
sensilla, tapering to a blunt point 47 behind level 
of sensillary centres* 

Crista surrounded by normal setae similar to 
those of whole of dorsum of idiosoma. Dorsal 
idioso-mal setae unexpanded, pointed, lightly 
setulose, curved, 18-46 long, the posterior setae the 
longer. There is no clear separation into 
morphologically different types of setae in any area 
of the dorsum. 

Ventral surface of idiosoma thickly covered with 
pointed, setulose setae, 15-34 long; seniles more 
prominent than for dorsal setae; setae less setulose 
but longer towards posterior pole, where they are 
similar to the posterior dorsal tdiosomalac. 

Genital aperture 86 long, by 50 wide; valves 
lightly chitinized; each medial valve with two or 
three setae 13-18 long at about mid-area, slender 
and less setulose than surrounding setae. Anus 49 
long by 31 wide, with 2 or 3 curved setulose setae 
along each lateral valve, similar to surrounding 


500 1 


i Wktf '\>\ ! --;. 

■ i -■■■'■ : \'AV- 

htg. 4 Odontacarui adeiaideae (Womersley). Deutonymph. 
entire, dorsal aspect, specimen ACB521AA/DJ. 

Legs long, fairly ihin; femoral to tibial segments 
more or less cylindrical lengths (including coxae 
and claws) I 960, II 570, 111 595, IV 925, Leg 
scobalae pointed, setulose; interspersed among 
them are many pointed sensory setae (spinalaej. A 
small vestigiala, 5-6 long, present on each of tibia 
I and genua I and II, distally and dorsolateral^'. 
Tarsus I outline and elongate oval, other tarsi more 
cylindrical (see fig. 4)- 

Other morphometric data as in Table 2. 

Palpi normal, with generally lightly setulose setae 
(many appearing aJmost nude), but along dorsal 
(extensor) edges of segments a number of 
unilaterally setulose setae (pectinalac). Palpal tibial 
claw stout, blunted, with four accessory thickened 
setae on dorso-medial aspect. T'alpal tarsus clavate, 
ovoid, with several setulose setae, but carrying also 
a cluster of terminal smooth sensory setae. 

Cheliceral fangs fairly robust, curved, blunt- 
pointed, with a row of about 10 minute denticles 
along flexor (concave) edge. 




Fig, J OdorUacarus udelaideae (Womerslev). Deutonvmph: gnathosoma and adjacent structures, dorsal aspect (partly 
in transparency), from specimen ACB521AA/U1. 


Table 2. Morphometry data for deutonymphs of Odontacarus adelaideae {Womersley). 












+ 12.71 






± 3.61 






± 3.83? 














Ta[(L) 3 










± 4.45 






± 5.58 






+ 10.51 





+ 13.27 






± 3.43 






± 8J1 


Till I 






Talll(L) 3 










+ 3.78 






+ 11.65 






+ 24,23 






+ 14.79 






+ 4.17 






± 0.056 


1 Cl_ = crisial length, measured from anterior Up of sagitta to level of midpoints of sensillary sockets 

2 For maximum values 

a Omitting claws and pedicle. 

Identification ol larvae of Odontacarus (Ifogonius) 
in Australia 


The recording of about 42 ventral opisthosomal 
setae in O. adelaideae larva invalidates caption 
number 12 of my earlier key to the larvae (Southcott 
1986a pp. 179-180), which was based largely on 
previously published descriptions. 

Accordingly, captions 12-14 should be replaced 
by the following: 

12 More than 45 ventral opisthosomaJ setae present . 

I ess ihan 45 (about 42) ventral opisihosomal setae 
present. AL seutalae in range 33-40 /xm lone. PDS 
34-46 ixm long O. adelaideae (Womcrslcy) 

13 More than 75 dorsal idiosonial setae present 

O. ouwalienste (Hirst) (including O. himti (Womersley, 


Less than 70 dorsal idiosomal setae present 14 

14 Pl./AL <1.20 O. novagumea 

(Womersley, 1944)3 (including O. longipes (Womersley. 
1945) 3 ) 

PL/AL>I.20 ... 


Key to deuionymph utstars of Lcogonius 
{Australiu-SW Pacific area) 

1 Dorsal idiosomalae arising fmm small platelets and 
about 25 (im long 1 

a, O. novaguinea (Womersley, 1944), 

Dorsal idiosomalae without the above combination of 
characters . , u . j 

2 Postcrioi dorsal idiosomalae over 50 /mi long, Tal/TiL 
mean 1.54 (tarsus I 210-228 ptti long, tibia I 130-154 
f*m) 2 O, austraiiensis (Hirst, 1925) 

(syu. O hirsti (Womersley, 1944)-V 
Postcriot dorsal idiosomalae generally less then 50p,m 
long _ H 3 

3 Tarsus I longer than 300 /tin (tarsus I 450 /mi, tibia 
1 450 /mi. Tal/Tii - 1.00, dorsal idiosomalae to 30 

Win long) 1 O. longipes (Womersley, 1945) 

Tarsus 1 less than 300 /nn long 4 

4 Posterior dorsal idiosomalae 15-20 /mi long, lal/Til 
1.36 (tarsus I 185 fim long, tibia I 136 ttm) 4 

O. uudyi (Radford, 1946) 

Postcrioi dorsal idiosomalae generally to about 45 /mi 
long. TaI(L)/TiI 1.15-1,30 (tarsus I 2*18-273 funlong, 
tibia I 178-226 /an O. adelaideae 

(Womerslev, 1944) 

i from Golf (1079). 

^ Synonymized by Domrow (1956) and Goff (1979). 

3 These two species (as larvae) were synonymized by 

Gof'f (1979), However, sec the comment of 

Southcott (1986a, p. 191). 

1 Womersley (1945) 

2 Domrow (1956) 
a GofF (1979) 

4 Nadchatrarn (1963) 


Fig. 6 Orion tacaritx adelatctMf (Wom&^Jey). Deuiortymph, 
external genitalia and anuii. from specimen 

Material examined r>fOdontacaru.s adelaideae: TVpe 
series: South Australia: Glen Osmond, Nov. 1931, 
D. C. Swan, from ears of cat, 3 paralcctotypes 
{NI98S333-NI98S335. ACU941-943). Wnfey Park 
(not Unley, as slateil by Womersley 1944), l.ii.1941, 
R V. Suuthcott. from ears of eat, one lectotvpe 
(NIV8833I . ACB67), one paralectotype (NI988332, 

The lectotype was. designated as NI988331 
because it has not been remounted since 1943 
indicated by my noresi and is. Therefore, the most 
intact Also it is in the same attitude as Womersley 's 
U944) figuie. 

f have excluded from the type series the three 
specimens recorded in the original description, from 
X'auns, Queensland, 1939 W. G. H^easbpV'from 
ra?s, n as tbey can no longer be found in the 
coUection of the South Australian Museum, and 
were presumably excluded by Womersley at some 
time after his 1944 paper. He did not reler lo them 
in 1945. \n his last publication on Qdortfoatrus, 
Other material: S. Aust. Unlcy Park (all from can 
of cats of topotype area, collected R.V.S.) l.iU94i, 

one specimen. ACB6H; 9.ii.l94», several specimen*. 
ACB72. 74 ? 928; l.iii.1941, 9, ACB75-77; ll.ui.194l. 
2, ACB 78.79, I5.iii.1941, 2, ACB80, 8J: 29.iii.l94l, 
U ACB82; 30.iii.194l, I, ACB83; 4.ivJ94|, I. 
ACB84; 6,iv.l94L L. ACB85; li.iv.194I. 3, ACB86 
B.C.E; i4.iv.1941, J, ACB87; 16.xi.1941. 3. 
ACBKXM02; 23.xi.l94L, 5, ACBI03: 29.xi.IWI, 5, 
ACB104; 22xiU946, several, ACB270; 16.xii.1947, 
several, ACB3I9; 3l.xii.1950, several, ACB520; 
31.xii.l950„ two batches of several larvae, from 
which deutonymphs were reared, ACB52I, 522; 
25.ii.l951, several, ACB523; l.iii.1951, several, 
ACB524; 9,iii.l95i, 2, ACB525; 6.1.1952. several, 
ACB560 (deutonymphs reared). 
Successful larva to nymph rearing experiementt 

Larvae were handled in the 1950-1952 scries of 
experiments- as stated above. One pmtunymph 
(ACB522BA5I was obtained, but the transformation 
was not observed, and discovered only when ihe 
tube was examined un 29X1951. Earlier examination 
on I4.U95I revealed nothing unusual (however in 
these wet tube studies observation is at times 

Eleven deutonyutphs were obtained from the 
1950-1952 experiment*. Nine of these came from 
experiments ACB521 aitd 522. with larvae collected 
on 31.xit.l9S0. and nymphs having emerged over 
15-29,1.1951. With batch ACB560, collected on 
6.1.1952, two newly emerged deutonymphs were 
found on 

The finding of an intact proconymph was 
lortunaie. as this instar i* generally unrecognizable 
after the emergence of the deumnvmphs- 

Kiom the above experiments, the larva to 
deuionymph transformations took 15-2V jfayj 
Domrow (1956) recorded that in O. austraiiensis 
deutonymphv look 25-28 days to emerge in his 
laboratory in Brisbane. 
Transfer of larvae of O. adelaideae from cats to man 

On the afternoon of 1 1 April 3941 I uans(emxl 
seven attached Jut vae (batch ACB86) from the ears 
of a cat to the skin of the dot sum of my forearm, 
under a watchglass kept tn place by sticking plaster. 
On removing the watchglass 2.5 hr later J found two 
small larvae attached to my skin. Removal of the 
watchglass periodically was necessary, as- 
condensation made detailed observations 

Next morning the underlying skin was sore. Three 
maculopapule.s with attached mites were observed 
at different sites; these maculupapules were 4-6 mm 
wide, and were itchy. That afternoon the area of 
erythema around the attachment sites had grown 
to 10-11 mm wide, these had become conjoined 
(only two mites were now attached). Small while 
specks were seen at the summih of I he papules (four 
beneath one mite, three beneath the other). 



presumably mite faeces. Small vesicles were present 
at the summit or each papule ai the mile tdtachment 
sites (20-22 hours after initial attachments (codes 
as S 4> 20-22 In)). 

The Itching, redness and vesicnlaUon increased. 
The attached mires QQ&cQ&rj slowly in size to "half- 
grown". At S + _*9 hr the whole area under the 
watchglass was itching, diffusely swollen and 
somewhat indurated; this area was 45 mm across, 
and by this stage the mites were considered full 
gmwn (as compared with (he larvae that had been 
observed on the cat's ears). The papules had become 
so prominent as to press upon the covering 
watchglass. The mites continued to make small 
faecal pellets beneath rhem. 

By S + c. $0 hr two vesicles had broken and were 
crying serum No lymphangitis or lymph- 
adenopathy had developed. 

After the miles had detached the swelling reduced 
rapidly, and scabs developed over the broken skin 

By S + 8 days the papules had retrogressed to 
being almost flat, and the erythema was lading No 
general symptoms attributable in the initc bitinj>s 
or their lesions developed, 

Unforiiinaicly, this attempt ro obtain mite 
dcutonymphs by human feedings was unsuccessful. 
A summarized account of this experiment was 
recorded earlier (Southcott, 1973). 


I thank the South Australian Museum for access 
to material in their collection, 

Tin's work was supported by the National Health 
and Medical Research Council and the Australian 
Biological Resources Study, 


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TrombicuiidatMAeaiiua), Mem, gfli. Soc. Washington 
4, [-W. 


Womersley, H. (1934) A revision of the trombid and Australia and New Guinea. Trans. R. Soc. S. Aust. 68 

erythraeid mites of Australia with descriptions of new (1). 82-112. 

genera and species. Rec. 5. Aust. Mus. 5 (2), 179-254. (1945) Acarina of Australia and New Guinea. The 

(1937) A revision of the Australian Trombidiidae family Leeuwenhoekiidae. Ibid. 69 (1), 96-113. 

(Acarina). Ibid. 6 (1), 75-100. — , & Heaslip, W. G. (1943) The Trombiculinae 

(1944) Notes on and additions to the (Acarina) or itch-mites of the Austro-Malayan and 

Trombiculinae and Leeuwenhoekiinae (Acarina) of Oriental Regions. Ibid. 67 (1), 68-142. 



by F. Tiver, A. D. Sparrow &R. T. Lange* 


The vegetation of inland north- western Eyre Peninsula, South Australia, is sampled for 
phytosociological analysis by transect quadratting. Using a combination of numerical classification 
(Bray-Curtis dissimilarity with average-linkage clustering) and ordination (detrended 
correspondence analysis), seven main vegetation associations are identified: Eucalyptus dutnosa- 
Westringia rigida, Eucalyptus gracilis-Stipa y Eucalyptus gracilis-Eucalyptus oleosa, Eucalyptus 
oleosa-Melaleuca pauperiflora, Eucalyptus oleosa-Enchylaena tomentosa, Eucalyptus brachycalyx 
and Eucalyptus yumbarrana-Triodia irritans. The last occurs on the siliceous Moornaba sands, and 
the other six on the calcareous Woorinen Formation. Environmental parameters correlate only 
weakly with a floristic gradation over the calcareous soils, suggesting that historical influences may 
be important in determining the distributions of the vegetation types. 

KEY WORDS: Eyre Peninsula, vegetation survey, vegetation analysis, classification, ordination, 
plant associations. 



by E Tiver. A. D. Sparrow & R, X Lange* 


Tiver, F. Sparrow, A. D. & Lance, R. T. ()989) The composition and distribution of the vegetation of 
north-west Eyre Peninsula. Trans. R. Soc. S. Aust. 113, 47-61, 31 May, 1989. 

The vegetation of iulaiid north-western Eyre Peninsula, South Australia, is sampled for phytosoetologieal 
analysis by transect quadratting. Using a combination of numerical classification (Bray-Curtis dissimilarity 
with average-linkage clustering) and ordination (detrended correspondence analysis), seven main vegetation 
associations aie identified: Eucalyptus dumosa-Westringia rigida. Eucalyptus gracilis-Sripti, Eucalyptus 
gratilis-Eucatyptus oleosa. Eucalyptus oleosa-Melaleuea pauperiflora, Eucalpytus oteosa-Enchylueno 
tomentosa. Eucalyptus brae hy calyx and Eucalyptus yumbarrana-Triodia irritans. The last occurs on the 
siliceous Moornaba sands, and the other six on the calcareous Woorinen Formation. Environmental 
parameters correlate only weakly with a floristic gradation over the calcareous soils, suggesting that historical 
influences may be important in determining the distributions of the vegetation types. 

Key Words: Eyre PemmMla,. vegetation survey, vegetation analysis, classification, ordination, plani 


Until recently, the vegetation of north-western 
T.yre Peninsula remained relatively unknown. The 
first detailed study was that of Crocker (J 946). 
Unfortunately the accompanying map only extends 
northwest as far as Poochera, where the vegetation 
is described as Eucalyptus oleosa - E.gracitis - 
Edumosa edaphic complex. Specbt (1972) also 
included only the southern-most portion of the area 
in his maps, describing the vegetation south of the 
Eyre Highway as Eucalyptus socialis - E. gracilis 
open scrub and that north of it as Eucalyptus 
incrassata - Melaleuca uncinata open scrub. 
Room s ma & Lewis (1980) described 16 
communities, defined by the tall dominants, which 
occur on north-west Eyre Peninsula. Their 
distribution maps arc not detailed enough to be able 
to extract the relative importance of any of these 

Descriptive surveys such as those mentioned 
above are not always capable of repetition by others 
(Nilsson 1986). For this reason, most modern 
surveys use quantitative data of the whole flora, and 
various forms of computer-assisted multivariate 
analysis, which result in a higher degree of 
objectivity and repeatability. A numerical 
classification of the vegetation of western Eyre 
Peninsula, one of the few such studies in South 
Australia, was undertaken by Margules 0c Nicholls 
(1987). Using complete species data from 104 plots 
each of 0.1 ha, cluster analysis was used to identify 
six vegetation communities on western Eyre 
Peninsula. This survey did not extend northwards 

onto the sand dune formations. Statistical models 
were constructed to relate the occurrence of the six 
communities to three environmental variables; 
depth of carbonate layer, distance from the coast 
and latitude Although allowing prediction of likely 
vegetal ion types in other remnant patches* these 
factors do not adequately describe the 
environmental factors important in determining I he 
vegetation pattern. 

The aims of this study were to classify the 
vegetation of north-western Eyre Peninsula (Fig. 1) 
using repeatable, numerical methods, and to provide 
some explanation of the main factors controlling 
the vegetation pattern. 

i ,..', .n .^H Port Augusia 

V'Dfi I iai:otn\«d. 

I i I 



* Dept of Botany, University of Adelaide, GPO Box 49*, 
Adelaide, S. Aust. 5001 

Fig. I. Showing the location of the study area on north- 
western Eyre Peninsula, SAust. 



Geology, Landform and Soils of the Study Area 

Most of the surface geological formations of 
western Eyre Peninsula (see Fig. 2) are derived from 
calcareous sands. It has been suggested that these 
sands were blown in from the exposed marine 
continental shelf during periods of low sea level and 
aridity, which occurred during the Pleistocene ice 
ages (Crocker 1946; Twidale & Campbell 1985). 
Another possibility is that the sands represent 
palaeo-dune systems that were formed by wave and 
wind action during periods of high relative sea level 
(Short ei at. 1986). 

The Bridgewater Limestone is composed of 
calcarenite t grains of calcareous sand cemented 
together with caJcitc crystals. In the maritime 2one 
it is exposed as coastal cliffs but is otherwise covered 
by unconsolidated, deep, calcareous shelly sand of 
low water-holding capacity (Wright 1985). The 
Bakara Calcrete consists of calcarenite in which the 
shell fragments have been progressively dissolved 
and replaced by nodular and platy calcrete. To 
reduce the ecological complexity of the survey, these 
coastal limestone formations were excluded from 
the study. 

The Woorinen Formation closely corresponds to 
the Chandada Plain landform in which the dunes 
have consolidated to form an undulating plain with 

no surface drainage (Twidale & Campbell 1985). 
Calcrete is usually present as a bed of nodules 
beneath the surface. The soils are calcareous 
throughout, and range in texture from sands to clay 
loams. They are usually shallow and reddish brown, 
but sandier grey varieties and redder varieties with 
higher clay content have been reported; they have 
been extensively cleared for agriculture (Wright 
1985). Areas of the calcareous sands normally 
associated with I he Bridgewater Limestone also 
occur on the Woorinen Formation. Near the coast 
these sands are coarse and shelly but further inland 
they are firmer with less-apparent shell fragments, 
Wright (1985) reports deficiencies in a wide range 
of minerals in these sands. 

The Molincaux or Moornaba Sand is of 
extremely recent origin (c 10 000 years b.p. onwards) 
and composed of sand dunes aligned in a NW-SE 
direction. Bleached white siliceous sands with a 
yellowish B hori?on occur on the dunes. The 
underlying soils of the inter-dune corridors contain 
calcareous earths typical of the Woorinen 

Sampling Strategy 

A system of transects was chosen as the best 
compromise between equal coverage of the region 

■ Cunfen* xj 

Fig. 2 The study area, showing major surface geological formations (after Rlisset 19fi9). 



Fig. 3 The study area, showing location of sites along seven irameeis. Shading represents uncleared vegetation. 

and efficiency of time spent on data collection. 
Transects were placed in suitable areas of native 
vegetation, running north and south from Ccduna, 
Maltee, Mudamuckla, Nunjikompita, Pimbaada, 
Wirrulla and Cungcna (Fig. 3). Transects were 
extended southwards only as fax as the boundaries 
of the Bridgewater formation and its overlying 
coastal sands, and northwards as far as was 
accessible by four-wheel-drive vehicle. In the 
northern part of the study area, sites were placed 
1 km apart, but in the southern part of the area, 
which is widely cleared for agriculture, native 
vegetation was often restricted to roadside 
remnants, and some sites had to be located as much 
as 2 km apart. 

Field experience showed that quadrats of 0.1 ha 
as recommended by Whiltaker (1978) and used by 
Margules & NichoUs f 1987) were an appropriate size 
lo achieve a good representation of species present 
without overlapping different vegetation types. One 
50 x20 m rectangular quadrat was measured at each 

Vegetation Data 

In each of the 162 sites sampled, aJI seed-plant 
specks were recorded, although only native 
perennials were used for analysis. Annual species 

were excluded because their occurrence depends on 
recent seasonal events; their inclusion would 
therefore lower the repeatability of the results. 
Introduced species were also excluded, since They 
do not form part of the natural vegetation pattern. 
Total cover was estimated for each species 
present, and expressed as a semi-quantitative score, 
an adaption of the cover scale as Braun-BLanquet 
(1932); see Table I. 

Environmental Data 

Anticipating some influence on flohstic 
composition by various disturbance factors- 
disturbance by erosion, grazing, animal tracks, 
undergrowth clearance and earthworks was visually 
assessed on a scale ranging from (nil) to 4 (severe). 
Sixty-two sites rated 2 (mild) or greater were 
excluded from analyses for definition of vegetation 

Landform was assessed in the field as dune-crest, 
dune-Hank, rise or plain (including wide interdune 
corridors). A soil core was dug in the centre of each 
quadrat and the depth to limestone recorded, in 
intervals of 10 cm. Texture was determined using 
the field method described by Northcote (1971). 
Existing rainfall maps were not detailed enough for 
the purposes of this study, so a rainfall map was 



constructed, using complete Bureau of Meteorology 
rainfall records for 24 recording stations within or 
near the study area. This enabled the average 
rainfall at each site to be estimated and classed into 
12 mm intervals. 
Vegetation Analysis 

The need to classify vegetation raises the question 
of whether vegetation types occur as discrete entities 
with distinct boundaries, or whether classifications 
are merely abstractions imposed by ecologists onto 
a continuously variable vegetation (Greig-Smith 
1983). If vegetation consisted of random 
combinations of species a continuum would result 
and classification into discrete groups would be an 
entirely arbitrary process. If, on the other hand, 
variation was always discontinuous and no 
intergradation occurred, there would be a finite 
number of vegetation types in existence, and 
classification would be a simple matter. The actual 
case lies between these two extremes, ranging from 
the near discontinuous and distinct vegetation types 
to almost continuous ecotones (Webb 1954). 

Consequently, a combination of two 
complementary analysis techniques is recommended 
when investigating vegetation patterns; a 
classification to define vegetation types, and an 
ordination to examine the relationships between the 
groups and the extent of ecotonal behaviour 
between them (Gauch 1982). 

For classification purposes, the dissimilarity 
between all pairs of sites was calculated with the 
Bray-Curtis distance coefficient, and then sites 

Tauix I. Semi-quantitative scores expressing percentage 
cover {adapted from Braun-Blanquet 1932). 

Cover Score 

Percentage Cover 

1 -2 
50 - 100 



6 5 
I — 

.70 .75 .80 .85 


| Etiei I y gt m dmnat a 

Wiitrin fl ii r ig I <l a 

Euctl> P .t_u_8 g racilis 
Sjipj sp. 


1 1 ) Eucalyptus a leg t a 

Eucaly ptus g racili 





| V Euci I y p t ua o I 9 o m a 

Mtlalsuca pau p Tlf Ion 

V F u c a l y p t us algan 

Encfijilaana tomentoiia 

V 1 E ueal^ptUB tt lt fiytily I 

V 1 1 Eucalyptus y umtaarrtna I A 

T r I a d i « irritant 

Fig. 4 Summaiy of dendrogram from clustering analysis showing arrangement of groups at 0.65 distance level. 



grouped from the least to the most dissimilar by 
the unweighted pair group method using arthmetic 
averages (UPGMA; Legendre& Legendre 1983) to 
produce a hierarchical dendrogram, the branches 
of which represent vegetation types. The 
complementary ordination method chosen was 
iktrended correspondence analysis, DCA (Gauch 
1982), Ordination methods such as DCA rank sites 
along a series of axes which describe the major 
floristic variation. The results are displayed as a 
scattergram from which continuity and disjunction 
of vegetation types can be determined and with 
which environmental factors can be correlated. The 
numerical taxonomy computer software package 
NTP (Belbin ei at. 1984) was used for both analyses. 
For the purpose of preparing vegetation maps, 
the sixty-two disturbed sites were returned to the 
data set, and a further cluster analysis performed. 
By the proximity of these sites on the dendrogram 
to sites of known vegetation type, most of the 
remaining sites were identified as belonging to one 
of the major vegetation types. 


The classification dendrogram is summarised in 
Fig. 4. Field notes indicated that the groupings 
defined at the 0.65 distance level were the most 
readily interpreted, a higher level producing groups 
with large internal variation, and a lower level 
producing too many groups for consideration. 
Seven major vegetation types were identified at the 
U.65 level, one of which, type VII, was widely 

dissimilar from (he other groups, only fusing with 
them at 0.90 level. A minor vegetation type was also 
identified. Represented by only one site, Cungena 
29, this site was dominated by Callitris preissii, 
rather than a eucalypt species. 

The relationships between the vegetation types 
can be seen from the ordination; a plot of the first 
two ordination axes is given in Fig_ 5. The first axis 
explained 50% of the variation in the data set, and 
corresponds to the first major branching of the 
dendrogram, separating vegetation type VJI from 
the ol hers, The separation between the remaining 
six communities is largely supported by the 
ordination, although there is a certain degree of 
overlap, most marked between types I and II, and 
types V and VI. The relationship of these six 
vegetation types to each other obviously teikds 
towards the "ecotone" rather than the distinct 
"association" end of the scale in vegetation pattern 
as described by Webb (1954). 

The species consistent in each of the vegetation 
types are presented in Table 2 and Fig. 6 summarises 
the associated landforms, soils and rainfall. Table 
3 gives a full listing of species found in the seven 
vegetation types. Understorey species arc equally 
important as shrubs and trees in delineating 
vegetation type. 

The vegetation types, their distributions (Fig, 7) 
and their environmental correlates are discussed 
below. The types have been named "associations' ' 
according to usual ecological practice, even though 
they do not fully fit the formal definition of 
"association" given by Braun-Blanquet (1932). 

AMIS I < V., i. li-iN t... >U,r, a .1 50%) 

Bg. 5 DCA ordination showing relationships between the seven major vegetation type;;. Symbols denoting vegetation 
type arc a* follows: O , I; A, II: 4 III; ■ IV; •, V; D. VI; Y, VIL 



Tabi r 2. Plant species characteristic of the seven major vegetation types, Marked species had u cover score of two 

or greater when present, and could he considered dominants. 



Key Species (75-100% of sites) Usually Present (50-74'ft of sires) 






Cassia nernophila 

var. nernophila • 
Eucalyptus dumnsa • 
Geijera linearifolia • 
Westringia ngida * 
Acacia coltettoides 
Eremophila glabra 
Melaleuca lanceolate 
Zygophyllum apiculatum 

Eucalyptus gracilis * 
Stipa sp. • 
Enchylaena tomentosa 

Geijera linearifolia 

Eucalyptus gracilis • 
Eucalyptus oleosa • 
Westringia rigida • 

Zygophyllum aurantiacum 

Eucalyptus oleosa • 
Melaleuca pauperiflora * 

Eucalyptus oleosa • 
Enchylaena tomentosa * 
Geijera linearijolia 
Zygophyllum aurantiacum 

Eucalyptus hrachycalyx 
Maireana erioclada 
Rhagodia crassifolia 

Eucalyptus yumharrana • 
Melaleuca eleuthrosiachya 
Triodta trritans • 

Eucalyptus gracilis • 
Eucalyptus oleosa • 
Diane/la revoluta 
Exocurpos aphyllus 
Pittosporum phyiliraeoides 
Rhagodia preissti 

Acacia colletioides • 
Eucalyptus oleosa • 
Eremophila glabra 
Melaleuca ianceolatu 
Sclerolaena obliquicuspis 

Eremophila scoparia • 
Triodia irr'uans • 
Maireana erioclada 
Hhagodia crassifolia 

Rhagodia crassifolia 

Maireana erioclada • 
Rhagodia crassifolia • 
Scteroleana diucanthu 
Sclerolaena obliquicuspis 
Threlkeldia diffusa 

Melaleuca pauperiflom • 
Enchylaena tomentosa 
Sclerolaena dtacanlha 
Threlkeldia diffusa 
Zygophyllum aurantiacum 

Dianella revoluta 

Eremophila crassifolia 
Podolepis capillaris 

The Vegetation Associations 

Association I: Eucalyptus dumosaAVestringia rigida 
The closest counterpart in the literature is the 
Melaleuca lanceolata association of Boomsma & 
Lewis (1972), which is said to occur with a number 
of species as co-dominants, including Eucalyptus 
dutnosa and E'. gracilis. Their classification includes 
a much wider range of vegetation associations than 
is here defined. It occupies a somewhat irregular 
distribution throughout the study area, restricted 
to rising ground on shallow calcrete, and the least 
sandy soils (sandy loams and loams). 

Association IF: Eucalyptus gracilis-Stipa 

This vegetation association could be described 
as a depauperate variation of Association I. 
Although some of the same species are present. 
Eucalyptus gracilis is the most important tall species 
uniting the group. The lower diversity understorey 

often includes a high percentage o( perennial grass 
{Stipa), The relationship of Associations I and II 
is indicated by their proximity on the dendrogram 
(Fig. 4) and overlap on the ordination (Fig. 5). 

The association has not been previously recorded 
on western Eyre Peninsula, and E. gracilis has only 
ever been reported as a co-dominant with other 
mallee species for the whole state (Boomsma &. 
Lewis. 1980). Its distribution is also irregular, 
occurring on the dry central to northern section of 
the Cungena transect, but also scattered through 
the southern Pimbaada and W'irrulla transects at 
higher rainfall, Very shallow, light soils on low lying 
ground are consistent with its occurrence. The 
resulting harsh moisture conditions probably reduce 
the number of understorey species present to those 
which can tolerate them, such as the shallow-rooted 
Stipa and Sclerolaena spp. 



Association III: Eucalyptus gracilis-oieosa 

Association III probably Forms the core of the 
Eucaiypius oleosa / E. gracilis association identified 
by Crocker (1946) and Spechi (1972), although 
Specbt appears to misidentify £. oleosa as £. 
soclalls. It is the most common vegetation 
association in the northern part of 1he study area, 
bul was not recorded by Margules & Nicholls (19K7). 
lis occurrence in the southern section of the 
Nunjikompita transect at slightly higher rainfall 
seems to suggest that its distribution is more 
restricted by soil than climatic factors. It is not 
present it; the south-western pari of the study area 
because the calcareous sands there favour other 
vegetation associations such as IV and VI, or on 
the south-eastern section where calerete outcrop* 
Favour Associations I and JI. 

The occasional occurrence of Association 111 on 
dune Hanks may be explained by historical factors. 
It is the most common inter-dune vegetation of the 
northern pari ol the study area* occurring on 
medium depth to shallow soils. Since the Molineaux 
sands have been deposited over only ihe last lew 
thousand years, they ate relatively unstable. In some 
instances sand from these dunes has drifted on to 
calcareous eaiths [II the mtcrdunes already occupied 
ly Axsociation 111 vegetation, and the long-lived 
species such as EL oleosa and K gracilis persist on 
3 soil which would perhaps noi at present favoui 
their establishment. The understotey in these eases 
tends to include species less typical of the 
association, such as Triodta irri/ans. 

Association IV: Eucalyptus oleosa-Metaleuca 
patiperi flora 

Association IV contains a very species-poor and 
sparse understorey of chenopods such as Rhagodia 
c/ussifofia and Matreana eriocluda, A known 
vegetation association including Eucalypti's oleosa, 
Rhagodia crassifolia. Zygophyllum auranuacum, 
Scteroiaetia diacantha and Melaleuca lanceoktta 
(Macules & Nicholls 1987), is probably equivalent 
lo it, their Melaleuca iunceolata a possihte 
misidentiftcation of M. pauper(lhra. 

Association IV occurs mainly in the 
\outhernwesiern part of the study area, on shelly 
calcareous sands, with scattered occurrences in (he 
north, for example, on the Nunjikompita transect, 
where outliers of this soil type occur on rises in the 
inter-dunes. These outliers arc probably lemnatus 
of the old coastal dune environment which would 
have been present before the Molineaux sands were 
deposited. Although t!^e vegetation association 
appears to occur at slightly higher rainfall, this may 
simply be a result of the soil type which favours 
its occurrence being chaiactcnsiic of the southern 
coastal higher rainfall areas. The low diversity of 
the association may be an indication that only a 

few- undeistorey species are able lo persist on Ihe 
coarse shelly sands. 

Association Vi Eucalyptus oleosa- F-ncbylaena 

Vegetation falling within Association V ts very 
variable and a large number of disturbed roadside 
sites were most closely allied ro u, indicating thai 
it may not be a naturally occurring association, but 
a result of disturbance. Many of the species are 
unpalatable, such as Geijera lineari 'folia, or low 
growing chenopods. which suggests removal of 
more palatable understorey species by grazing. 
Further investigation would be required to ascertain 
if any of the sites represent a naturally occurring 
Association V, 

Association VI. fcuealyptus brachycalyv 

Vegetation Association VI includes Eucalvfitu* 
brachycatyx as the important upper-slorey species, 
together with a range of low chenopods similar to 
thai of Association V. The floristic similarity result; 
in considerable overlap between these two 
associations on the ordination <Hg. 5), (l is most 
common towards the coast, occurring in the 
southern central and western part of the study area 
on shallower soils than Association IV 

A\sociaiionV!i; Eucalyptus yumbanana-THodia 

it is probable that hucuiyptus yumbarrana was 
previously recorded as Eucalyptus sociafis, and that 
the E. soaalis/ Melaleuca uncinata sandhill 
association pi Specbt (1972) corresponds to 
Association VII, as docs Macules & Nicholls* 
Community I il their Melaleuca adnata is re- 
idetuifled as M eleuiherostuchya. 

Vegetation Association VII is restricted to the 
sand-dunes of the northern, low rainfall part of the 
study area. The speclflS characterising it are entirety 
different from those found in the other vegetation 
associations, and include many species with 
distributions in higher rainfall parts of the state, 
such as Lomandra leucocephah and Gaknia 
ianigera (Jessop & Toelken 1986). This major 
flonstic disjunction corresponds to a similar 
observation in north-western Victoria, where the 
most marked disooiuinuitv of tnailec vegetation was 
between a Eucatyp/iti iturossata/Hr'bbertia/ 
Leptospennum association with ''southern 
temperate" affinities, and all the other associations, 
»n which semi-succulent herbs, especially 
chenopods, were prominent (Noy-Meir 1971). 

The disjunction observed is related to Uie geology, 
since the heavier soils tS the exposed Wooritwm 
Formation present in the inier-dune corridors 
!>picaJJy support vegetation of Associations i, H 
ot Oil The combined effect of coaise texture ami 







p r f a 

20 4C 6p > ?D 


t 3L LS S 

Fig 6. Histograms summarising observed environmental patterns for each of the vegetation types. For topography: 
P, plain; R, rise; F, dune flank; S, sandhill. For soil texture: L, loam; SL, .sandy loam; IS, loamy sand; S, sand, 

increased depth of the sands of the dunes 
presumably creates a more favourable water 
relations environment, which allows the presence 
of a suite of species normally requiring higher 
rainfall There is a higher percolation rate and 
deeper penetration of rainfall into sands than into 
clays, while evaporation dries the surface soil to a 
similar depth regardless of texture, so that at the 
end of a long dry period, there would be more water 
available in the root zone of sandy soils than in clays 
(Walter & Stadelmann 1974). Clays also have a 
higher water holding capacity due to their smaller 
pore size (Ball 1986). This means that a larger 
amount of rainfall is required to bring air-dry clay 
to a range where water is available to plants than 
for sands. 


There is a clear vegetational discontinuity 
between the two predominating geological 

formations of north-western Eyre Peninsula, the 
siliceous Moornaba sand dunes and the calcareous 
Woorinen soils. Although it is possible to recognise 
plant associations within the vegetation of the 
Woorinen Formation and to relate their 
distributions to environmental features in a general 
way, sharp discontinuities do not exist. The strong 
gradation of associations is typical of the mallee 
on Woorinen Formation and its equivalents across 
the whole of S. Aust. (Sparrow unpublished data). 
There are two possible interpretations of these 
observations. Firstly, since there is no strong 
environmental discontinuity within the Woorinen 
Formation, the gradation may reflect combinations 
of species with independent environmental 
responses, which, further confounded by 
disturbance, results in almost continuous ecotones. 
Alternatively, the gradation of mallee associations 
could be caused by some degree of independence 
of the malice and understorey slTata. The mallees, 



Fig. 7 Distribution maps of the seven major vegetation types. Shading represents uncleared vegetation and the broken 
line marks the Eyre Highway. 



Table 3: Full species list {nomenclature following Jessop & Toelken 1986) showing occurrence in the seven major 
vegetation types. Occurrence in sites that could not be assigned to a major vegetation type are grouped in the final 
column. Introduced species are indicated with an asterisk. 


Vegetation Type 



VII Other 

Acacia calamifolia 

Sweet ex Lindley 
Acacia colletioides 

Acacia hakeoides 

Cunn. ex Benth. 
Acacia ligulata 

Cunn. ex Benth. 
Acacia merralli 

F. Mucll. 
Acacia notabilis 

F. Muell. 
Acacia oswald ii 

E Muell. 
Acacia rigens 

Cunn. ex Don 
Acacia sclerophylla 

Acacia spinescens 

Alyxia buxifolia 

R. Br 5 
Atnyema melaleucae 

(Miq.) Tieghem 
Atriplex acutibractea 

R. Anderson 
Atriplex stipitata 

Atriplex vesicaria 

Heward ex Benth. 
Beyeria lechenaultii 

(DC.) Baillon 
Beyeria opaca 

F. Muell. 
Biltardiera cymosa 

F. Muell. 
Boronia coerulescens 

F. Muell. 
Cailitris preissii 

ssp. preissii 

Cailitris preissi 

ssp. verrucosa 
(Cunn. ex Endl.) .1. Garden 
Carpobrotus rossii 

(Haw.) Schwantes 
*Carrichtera annua 

<L.) DC. 
Cassia nemophila var coriacea 

(Benth.) Symon 
Cassia nemophila var nemophila 

Cunn. ex J. Vogel 
Cassia nemophila var platypoda 

(R. Br.) Benth. 
Cassytha glabella 

R. Br. 
Cassytha melantha 

R, Br. 
Casuahna cristata 

Chenopodium desertorum 

(J. Black) J. Black 
Cratystylis conocephala 

(E Muell.) S. Moore 

+ + + 

+ - + 



t 4 



Table 3 cont. Full species list (nomenclature following Jessop & Toelken 1986) showing occurence in the seven 
major vegetation types. Occurrence in sites that could not be assigned to a major vegetation type are grouped 
in the final column. Introduced species are indicated with an asterisk. 


Vegetation Type 





Cryptandra leucophracta 

Cryptandra tomentosa 

Danihonia spp. 

Lam. & DC. 
Daviesia asperula 

Daviesia benthamii 

Dianella revoluta 

R. Br. 
Dillwynia uncinata 

(Turcz.) J. Black 
Dodonaea bursariifolia 

F. Muell. 
Dodonaea stenozyga 

F. Muell. 
Dodonaea viscosa 

ssp. angustissima 
(DC.) J.G. West 
Enchylaena tomentosa 

R. Br. 
Eremophila crassifolia 

(F. Muell.) K Muell. 
Eremophila deserti 

(Cunn. ex Benth.) Chinn 
Eremophila glabra 

(R. Br.) Ostenf. 
Eremophila scoparia 

(R. Br.) F. Muell. 
Eremophila weldii 

F. Muell. 
Eucalyptus brachy calyx /concinna 

Blakely/Maiden & Blakely 
Eucalyptus dumosa 

Cunn. ex Schauer 
Eucalyptus foecunda 

Eucalyptus gracilis 

F. Muell. 
Eucalyptus incrassata 

Eucalyptus oleosa 

F. Muell. ex Miq. 
Eucalyptus sociaiis 

F. Muell. ex Miq 
Eucalyptus striaticalyx 

W. FiLzg. 
Eucalyptus yumbarrana 

Eulaxia microphylla 

(R Br. in W.T. Aiton) J. Black 
Exocarpos aphyllus 

R. Br. 
Exocarpos sparteus 
R. Br. 
Gahnia lanigera 

(R. Br.) Benth. 
Geijera linearifolia 

(DC.) J. Black 
Goodenia varia 

R. Br. 

+ + + + 

+ + +■ + + 

+ -+■ + f t- 

-i- + + + + 

+ I + + + 



Table 3 cont. Full species list (nomenclature following Jessop & Toelken J986) showing occurence in the seven 
major vegetation types. Occurrence in sites that could not be assigned to a major vegetation type are grouped 
in the final column. Introduced species are indicated with an asterisk. 


Vegetation Type 





Grammosolen truncatus 

(Ising) Haegi 
Grevillea huegelii 

Hakea francisiana 

Ualgania cyanea 

Halgania tavandulacea 

Helichrysum bilobum 

Hybanthus floribundus 

(Lindley) F, Muell. 
Ixiolaena pluriseta 

Lasiopetalum behrii 

K Muell. 
Lepidosperma laterale 

R. Br. 
Leptospermum coriaceum 

(K Muell.) Cheel 
Leucopogon cordifoiius 

Logania nuda 

F. Muell. 
Lomandra collina 

(R. Br.) Ewart 
Lomandra effusa 

(Lindley) Ewart 
Lomandra leucocephata 

(R. Br.) Ewart 
*Lycium ferocissimum 

Maireana appressa 

Paul G. Wilson 
Maireana brevifolia 

(R. Br,) Paul G. Wilson 
Maireana enchylaenoides 

(F. Muell.) Paul Ci. Wilson 
Maireana erioclada 

(Benth.) Paul G. Wilson 
Maireana pentatropis 

(Tate) Paul G. Wilson 
Maireana sedifofia 

(F. Muell.) Paul G. Wilson 
Maireana trichoptera 

(J. Black) Paul G. Wilson 
Melaleuca acuminata 

F. Muell. 
Melaleuca eleutherostachva 

F. Muell. 
Melaleuca lanceotata 

Melaleuca pauperiflom 

F. Meull. 
Microcybe multiflora 

Muehlenbeckia gunnii 

(Hook, f.) Walp.) 
Myoporum platvcarpum 

R. Br. 
Ofearia brachyphylla 

(F. Muell. ex Sonder) Wakef. 

+ + 


* f 



Table 3 cont. Full species list (nomenclature following Jessop & Toelken 1986) showing occurence in the seven 
major vegetation types. Occurrence in sites that could not be assigned to a major vegetation type are grouped 
in the final column. Introduced species are indicated with an asterisk. 


Vegetation Type 



vi r 


Olearia floribunda 

(Hook, f.) Benth. 
Olearia magniflora 

(E Mucll.) F. Mucll. ex Benth.) 
Olearia mue/leri 

(Sonder) Benth. 
Olearia pimelioides 

(DC.) Benth. 
Pimelea microcephala 

R. Br. 
Pittosporum phylliraeoides 

Podolepis capillaris 

(Steetz) Diels 
Ptilotus obovatus 

(Gaudich.) E Muell. 
Ptilotus spathulatus 

(R. Br.) Poiret 
Pultenaea elachista 

(E Muell.) Crisp 
Rhagodia crasstfolia 

R. Br. 
Rhagodia parabolica 

R. Br. 
Rhagodia preissii 

Rhagodia spinescens 

var. deltophylla 

R Muell. 
*Salsola kali 

Sanialum acuminatum 

(R. Br.) A. DC. 
Scaevola bursariifolia 

J. Black 
Scaevola spinescens 

R. Br. 
Schoenus subaphyllus 

Sclerolaena diacantha 

(Nees) Benth. 
Sclerolaena obliquicuspis 

(R. Anderson) Ulbr. 
Sclerolaena uniflora 

R. Br. 
Sida corrugata 

*Solanum coaciiliferum 

J. Black 
Stelligera endecaspinis 

A. J. Scott 
Stipa spp. 

Templetonia battii 

F. Muell. 
Threlkeldia diffusa 

R. Br. 
Triodia irritans 

R. Br. 
Velteia connata 

E Muell 
Vittadinia dissecta 

(Benth.) N. Burb. 

+ + + + 

+ + 

+ + 

+ t 

+ + 

+ + 

+ + 


E T1VER, A. D. SPARROW &. K. I. | ANOfc 

Tahi p 3 coot. Full species list (nomenclature following Jessop & Tadkert IP86) showing occurence in the seven 
mq/or vegetation types. Occurrence in Sites that could not he aligned to a mnjor vegetation xvp& are grouped 
in the final column Introduced species arc indicated with an asterisk 


Vegetation type 

Htestringta hgida 

R. Br. 
Zygophvllum ammophUmt 

F Muell. 
ZvROphylhnn apicutatum 

F, Muell. 
Zygophytlunt uUrontiacum/ovatum 

iXindicy) F. Muell./twarl & .1. White 
Zygophvllum glaucum 

F, Muell. 


• I 

A + 





being very long-lived plants, may be indicating some 
kUiorical influence, while the shorter-lived 
understorey species reflect current environmental 
conditions. Lange &, Nicolson (1982) report mallees 
recording former extensions of a palaeosoi, and 
Twidalc & Campbell (1985) and Short el ai (I9K6) 
provide ample evidence of dynamic changes in the 
surface geology of norlh-western Byre Peninsula 
during the Holocene. 

A further issue is the occurrence of uncommon 
vegetation associations in the area. One example is 
the Caflitris prgjssfi association represented by a 
single site on the dendrogram; another, towards the 
northern end of the Mudamuckla transect and 

removed from analysts due to disturbance, was 
unique For the presence of Casuarina cristata. A 
regular sampling strategy inevitably leads to tfo 
omission of rare vegetation types, so (here are 
undoubtedly other uncommon associations in the 


We would like to thank the staff of the Vegetation 
Retention Branch of the S Aust. Department of 
Environment and Planning for practical assistance 
during the course of the study. The work was 
carried out as pai i of the Middleback Field Centre 
programme for 1987. 


Km i O. h (1986) Site and soils pp 215-284 In R l> 

Moore &. S.B. Chapman (Z,c\a) y "Method* in plant 

ecology" (Klackwcll Scientific Publications, Oxford). 
Hi i WN.1,,, Faith. D. ft it Mincmin, P. R. (1984) "Some 

algorithms contained in the Numerical 'taxonomy 

Package NTP" (CSIRO Division of Wato and Land 

Resources, Canbetra). 
Blissett.A. JL (1969)STRF.AKY HAY preliminary map 

sheet, Geological Survey of Sourh Australia 1:250 OCX) 

series (Department Of Mines and Energy, S. Aust.). 
Boomsma. C. 0. & Lewis, N. B. (1980) 'The native foreM 

and woodland vegetation of South Australia" (Woods 

and lorests Department, S. Aust.). 
Braun-BlaNQUEI. J. U9J2) "Plant sociology" (McCiraw- 

Hill, New York & London). 
Ckocklk, R. I.. (lV4o) An introduction to the soils and 

vegetation of Eyte Peninsula, South Australia. Trans. 

R. Sot: S. Aust. 70, 83-107. 
CtAitf.H, H. G. (1982) "Multivariate analysis in community 

•ecology" (Cambridge University Press, CamlvkJ^c). 
CrRRG-SvnTH. P (1983) "Ouanlitulive plant ecology" 

(Blackwcll Scientific Publications, London). 
Jessop, .1. P. & Toelken. U. R. (Eds) (1986) "Flora of 

South Australia" Parts I— TV (S. Aust. Govt. Priming 

Division, Adelaide). 
Lavjl, R. T. & Nitotsos, k. P. (19X2) Vegetation 

apparently recording fotmr.r extensions nf a palcosol. 

Trans. H, Soc. & Attsi. 106. 77-78. 
I.Kii'Nimfi, L, <fe LecLNDfte, P. (198*) "Numerical 

ecology" (Elsevier, Amsterdam). 

MAKGULES..C. R. £c Nicnor i s. A. O. (1987) Assessing the 
conservation value of lemnam habitat islands": Malice 
patches on the Western fcvre Peninsula, South Australia, 
pp 89—102 In t). A. Saunders, U. W. Arnold, A. A. 
Burbridge & A, J, M. Hopkins (P.ds) "Natutc 
Conservation: The Role of Remnants of Native 
Vegetation" (Surrey Beatty & Sons with CSIRO, Penh>. 

NlLSSON, L. (19S6) Method's Of >eleetiiu; lake shorelines 
as nature icsv'ves. Biol. Cons. 35, 269-2W. 

Northcote. K. R. (1971) "A lactual Key I'd the 
Recognition of Australian Soils" (CSJRO, South 

Nov Mi-jr. r, (J971) Multivariate analysis of the semi- 
aiid vegetation of south-eastern Austialia; nodal 
ordination by component analysis Proc. Ecol. Soc. Au.u. 
6, 159-193. 

Short. A. D., Fothekim.ham, D. G. & Buckley. R. C. 
(1986) "Morphodymimie, Hazaid and Development 
Impact Assessment - Eyre Coasi Protection District" 
(Department of Environment & Planning, Adelaide). 

Specht. R. L. (1*72) 'The Vegetation of Souih Australia" 
<S. Ausr. Govt, Printer. Adelaide). 

Twidm t-, C. R. <fc Campbell Z. M. [V$$) t\w J'oim of 
the land surface. In C. R. Tv»ndalc, V1. J. Tyler & M. 
Davie? (Eds) 'TMatural History ol Lyre Peninsula" (K. 
Soc. Si AuSt, 4 Adelaide). 

W'mter. H. ik. Stadelmann, L. (1974) A new approaeli 
to Hie water relations ordevrn plants, pp 213-310 /n 
G. W. HroHn tKd.) " Uioloj^y: Special Topievod 
the Physical and Biological Aspects of Arid Regions" 
I Academic PQtM, New York). 


Webb, D. A. (1954) "Is the classification of plant "Ordination of Plant Communities" (W. Junk, The 

communities either possible or desirable?" Botanisk Hague). 

Tidsskrift 51, 359-380. Wright, M. J. (1985) Soils, pp 77-87 In C. R. Twidale, 

Whittaker, R. H. & Gauch, H. G. (1978) Evaluation of M. J. Tyler & M. Davies (Eds) "Natural History of Eyre 

ordination techniques, pp 277-336 In R. H. Whittaker Peninsula" (R. Soc. S. Aust., Adelaide). 




VOL. 113, PART 2 





Stream erosion on the eastern flank of the Mt Lofty Ranges has exposed a reverse fault near 
Cambrai on the Milendella escarpment, indicating a compressive component to the uplift of the 
ranges. Cambrian rocks have been thrust over Pleistocene fanglomerates and Miocene limestone 
has been dragged up along the fault zone to an elevation of 160 m asl. This fault may indicate the 
reactivation of a compressive Palaeozoic fault during the Cainozoic. Diagnostic foraminifera have 
been identified in samples of the limestone, which is of the Early Miocene Mannum Formation 
(about 20 Ma), adding support to the unpublished view that the position of the limestone on the 
escarpment is due to tectonic uplift of 60-90 m since the Miocene. 

KEY WORDS: Tertiary limestone, foraminifera, reverse faulting, Mt Lofty Ranges, South 


by R. P. Bourman* & I M LindsayT 


Boukvias, K. P. & Lindsay, J. M. (1989) Timing, extent and character of late Cainozoic faulting on the 
eastern oiarcin of the Mt Lofty Ranges. South Australia. Trans. H. Soc. S. Aust. 1 13< ). 63 -67. 31 May. 1989. 
Stream erosion on the eastern flank p! (he Ml Lofty Ranges has exposed fi feverse taull near Cambrai 
on the Milendella escarpment, indicating b compressive component to the uplift of the ranges. Cambrian 
foek* have been thrust over Pleistocene umglomeratei and Miocene limestone has been dragged up along 
the fault zone to an elevation of 160 m n«i|. this ffttfll may indicate the reactivation of a compressive Palaeozoic 
fault during the Caiiio/oic. Diagnostic foraminifera have been identified in samples of rhe limestone, which 
is or rhe Early Miocene Mannum Formation (about 20 Ma), adding support to the unpublished view dial 
the poiiiioit of the limestone on the escarpment is due to tectonic uplift of 60-90 m since the Miocene. 

Knv Words Tertiary limestone, foraininifeia, reverse faulting. Mi Lofty Ranges, South Australia, 


The fault origin of the Ml Lolly Ranges, hux totlg 
been discussed (Benson 1911; Fenner 1910; Sprigg 
1945) and the majority of these workers considered 
that the faults were normal types. Glaessner (1953) 
presented a model for the tectonic evolution of the 
Ml Lofty Ranges, which involved basement complex 
fleixuring, that resulted in reverse faulting on the 
margins of the ranges and normal faulting within 
them. Campana (1955) preferred to explain the 
origin Of rhe Mt Lofty Ranges by cornpressional 
doming, which kd to minor fault disruption on 
Iheir margins, Gibson (1963 ) 1 noted that the 
Clarendon-Ochre Cove Fault displays a reverse habit 
where intersected by tunnelling operations during 
the construction Of l he Clarendon-Happy Valley 
pipeline. Recently Well man & Greenhalgh (1988) 
favoured the view that compressive forces have been 
important in the formation of the Mt Lofty Ranges, 
A rc-evalualion of exposed fault contacts is required 
to resolve the nature of the faulting responsible for 
the uplift of the Mt Lofty Ranges. 

The Milendella Fault 

There is clear evidence of reverse faulting on the 
eavtern escarpment of the Mt Lofty Ranges as first 
reported by Mills (I965) 2 , and it may indicate the 

' Gibson. A. A. (196?) Final geological repott on the 

Happy Valley 'funnels Project- S- Ausi. Dept of Mines 

rrport 57*92 funpubl.) 
- Mills, K. J. U%?> 'The structural petrology Of the 

Milendella area of South Australia". Ph.D. Thesij. 

University of Adelaide (unpubl.) 

* Depx of Soil Science, Wake Institute, university of 

Adelaide. Urrbrae, S. Aust., 5064. 
t i ivolo^icid Survrv, South Au^lralLin Dcp.irtiin n( ftl 

MtntS & r.ncrgy. Box 151, dust wood P.O., S. Ausi. 5063. 

re-activation of Palaeozoic ihruM faults. Stream 
exposures of the Milendella Fault occur west of 
Camtocal on the escarpment (Angaston MR 
390/630) at 160 m asl (Fig. 1). The fault here is of 
reverse type with brecciated Cambrian schists of the 
kanmauton Group of metasedimentary rocks thrust 
over Pleistocene fanglomerates, Tvvo pods of 
Tcrtiar) limestone have been dragged up at a high 
angle in the fault zone, which dips to the west al 
45° {Fig. 2). The Pleistocene fanglomerates affected 
hy the faulting are similar to PLeistocene 
fanglomerates tilted into <t vertical position by 
movement along the Willunga Fault as Selheks 
Reach (May & Bourman 1984). The sediments al 
Selticks Beach were related to the Ochre Cove 
Formation, which May «& Bourman (1984) regarded 
as of Middle Pleistocene age. If the two deposits 
are correlatives then faulting on the Milendella 
Scarp probably occurred until the late Middle 
Pleistocene oi the early Late Pleistocene. No 
unequivocal evidence of dislocation of Late 
Pleistocene sediments in the area was noted. 

It is suggested that during faulting* the 
Pleistocene fanglnmerate* were folded, and 
brecciated basement rocks broke from the hanging 
wall to tumble on top of the fanglomerates. Calcrete 
mantles the convoluted fanglomerates. dying out 
at the latih zone, so that the calcretc may have 
derived from solution and reprecipiiaiion of the 
Tertiary limestones dragged to the surface along the 
fault /one and draped over the top of the 
fanglomerates. Calcareous pisoliths in ihb calcrete 
are typically laminated around cores of locally 
derived bedrock. This supports the interpretation 
of in situ pisolkh formation from the redistribution 
of lertiary limestone. Alternatively, if the calcrete 
has stratigraphic significance (e.g. the upper Middle 
Pleistocene Bakara calcrete of Firman 1967) then 




^Y(V\ PMANNUM Teal¥Flat 

CM Walker Flat 

\ o(murray BRID 






Fig. I I miuiiiri map sliuwing major fault zones, ' site of fault exposure near Cambrai and a Leprdocyc/lnu-btM'm^ 
limestone in the Biemer valley. 

the relationships may indicate fault disruption of 
the calcrete in the Late Pleistocene. It is not possible 
to determine the amount of olTsel accomplished 
during this postulated Pleistocene phase of faulting 
without detailed drilling. However, Mills (1965) 2 
shows a thickness of some 30 m of Pleistocene 
sediments in section at the scarp (Fig. 3), and as 
the limestone must have been dragged up from 
below this level, this suggests an offset of less than 
30 m. Dislocation of Early Pleistocene marine 
sediments across the Willunga Fault was reported 
by May & Bourman (1984) illustrating similar post- 
Farly Pleistocene faulting of about 40 m on the 
opposite side of the ranges. It is interesting to note 

that although there is clear evidence of geologically 
recent faulting on the eastern margin of the Mt 
Lofty Ranges, there is no pronounced fault scarp 
associated with it, which may suggest that (he scarp 
formed largely in brecciated rock and rapidly 

Mills (1965)2 suggested thai displacement on the 
Milendella Fault since "p re -Tertiary peneplanation" 
was approximately 335-366 m, and he attributed 
250 m of this to the Early Tertiary to Miocene, and 
60-90 m to post-Miocene faulting. He noted 
Miocene sediments (?Morgan Limestone) at 
elevations up to 160 m on the scarp, correlated them 
with Miocene strata under the Murrav Surface and 




. ./. 

■ :*-";Hi"- *■-■%. : . 


Fig. 2. Fault exposure near Cambrai. View south along the eastern escarpment of the Mt Lofty Ranges. Cambrian 
schists (K) of the Kanmantoo Group of metasedimentary rocks have been thrust over (?) Middle Pleistocene 
fanglomerates (F). Light coloured calcrete (C) mantles the fanglomerates and two pods of Miocene limestone (L) 
occur in the fault zone (arrowed), which dips to the west at approximately 45°. 

attributed the difference in elevation to tectonic 
dislocation. Twidale& Bourne (1975) did not deny 
recent fault activity but preferred to interpret the 
perched limestone as essentially due to Tertiary 
eustatic influences, with the limestone under the 
Murray Basin being the older, lower Mannum 
Formation and that on the scarp as the younger and 
higher Morgan Limestone. Thus they discounted the 
view that there had been 60-90 m of dislocation 
on the Milendella Fault since the Miocene and 
argued for an erosional origin for the lower part 
of the scarp. The resolution of this conflict may be 
achieved only by an accurate age determination of 
the Miocene limestones on the Milendella scarp to 
compare with those from the Murray Basin. 
Palaeontological work reported here demonstrates 
that the limestone on the scarp, at least in this 
locality, is from the lower part of the Mannum 

Age and facies of the Tertiary limestone in the fault 

Four samples have been examined from the two 
pods of Tertiary limestone dragged up in the fault 

Portions were crushed gently in a pestle and 
mortar to disaggregate partially, boiled in a dilute 
solution of sodium bicarbonate with a drop of 
detergent, washed free of mud through a very fine 
sieve, dried, and picked for foraminifera under a 
binocular microscope. Slides with the foraminiferal 
microfaunas are stored in the Biostratigraphy 
Branch collections of the S. Aust. Department of 
Mines and Energy. 


All four samples comprise quartzose fossiliferous 
sandy limestone (biogenic calcarenite). The rocks 
are cream to brown, hard to friable, somewhat 
recrystallised and, in part, leached. Quartz and 
minor lithic sand content is considerable and poorly 
sorted, ranging from very fine grained to very coarse 
grained. Quartz and lithic grains up to 10 mm are 
present. There are occasional pale green glauconite 
infillings, somewhat oxidised. 

Barnacle plates are frequent and notable; 
echinoid, molluscan and bryozoal fragments are 
frequent to common; foraminifera are common but 
mostly recrystallised; ostracods, Ditrupa tubes, fish 
and decapod fragments, and algal oncolites are rare. 




a> 300 


% 200 



V.E. X20 


Pleistocene alluvium 

Displaced Miocene limestone blocks 

Recent alluvium 


Miocene limestone 


Eocene lignitic sands 

Fig. 3. Cross section through the eastern margin of the Mt Lofty Ranges to the Murray Basin near Cambrai (after 
Mills 1965 2 ). 

Discussion of foraminifera, age and correlation 

The four samples have much in common and the 
microfaunas will be considered together (Table 1). 
Numerically, all the microfaunas are dominated by 
Elphidium chapmani and Notorotalia cf. 
miocenica, but these are not the most 
biostratigraphically significant. 

The overlap of the top of the range of 
Sherbornina atkinsoni and the bottom of the range 
of Operculina victoriensis in S. Aust. indicates an 
early Longfordian, Early Miocene age (Wade 1964; 
Lindsay 1985). For the Murray Basin proper, 

Ludbrook (1961) only overlapped these ranges in 
lower Mannum Formation, and this has been borne 
out by subsequent experience. Other foraminifera 
present in the four samples are generally consistent 
with Longfordian Stage and Early Miocene age. 
Notorotalia miocenica has been scarcely recorded 
previously from S. Aust., so the slightly earlier age 
here of a related form, compared with the range 
in Victoria (Carter 1964) is not regarded as 
incongruous. The solitary and small planktonic 
specimen identified as Globigerina sp. is 

Table 1. Foraminifera: Table of occurrences of selected species. 



F24/83 F2/88 


Discorbis sp. from group D. balcombensis 

Chapman, Parr & Collins -D. cycloclypeus 

Howchin & Parr 

Ammonia sp. 

Pararotalia verricutata (Howchin & Parr) 

Sherbornina atkinsoni Chapman 

Elphidium chapmani Cushman 

E. crassatum Cushman 

E. parri Cushman 

Notorotalia cf. miocenica (Cushman) 

Operculina victoriensis (Chapman & Parr) 

Globigerina sp. 

Amphistegina sp. 

Cibicides vortex Dorreen 

Planolinderina plana (Heron-Allen & Earland) 





















































(Abundances: V = very rare; only one or two specimens recovered in thorough picking. R = 
6-10. C = common; 11-25. A = abundant; more than 25). 

rare; 3-5. F = frequent; 



The liihology of (he samples is consistent with, 
bui not unique to, Mannum Formation as described 
by Ludbrook (1961) and observed subsequently. This 
tarly Miocene Mannum Formation in the fault 
*one is distinctly older ihan the elevated, 
Let)tdocyiiiri<j-be<\£, Morgan limestone reported 
!>om the Bremer valley east of Kanmamoo (Lindsay 
I986 1 ); about 20 Ma compared with 16 Ma (Lindsay 
J 985). 

tnvirortment of deposition 

The variety and nature of the fauna indicate 
marine conditions. Three lines of evidence suggest 
a shallow-marine, liuoral-neritic environment: the 
notable frequency of barnacle remains, the 
restricted Eiphidium-r'\cU foratniniferat fades 
almost lacking planklonic forms, and the quanzosc 
and lithic clastic component of the limestone, with 
gravcl-si/ed grains, This corroborates the 
scdimcntological evidence of Mills (l%5i^, who 
proposed a near-shore unvironmem fnr i.he 
deposition of (he limestone. 


The evidence presented here demonstrates thai 
most ol the elevalional difference between the 

Mannum Formation limestones on I he escarpment. 
near Cambrai, and under the Murray Basin can be 
aitributed to tectonic dislocation in the past 20 Ma 
as- originally suggested by Mills <J965) ? . 
Furthermore, it has also been demonstrated thai 
i here 1ms been Jit least I IK) ffl (and probably much 
more) of tectonic offsei nf the early Middle- 
Miocene Upkk>tyditw~bcai'\i\g Morgan Limestone 
that occurs in the Bremer Vailey (I indsay I986) 3 
in the pasi 16 Ma. Consequently, evidence of 
considerable tectonic dislocation of Tertiary 
limestones on the eastern side of the Mr Lofty 
Ranges, separated by 50 km, together wuli faulted 
Pleistocene sediments provides strong support for 
pronounced tectonism since the Middle Miocene. 
Without denying the evidence lor euslalk: influences 
in the Tertiary {e.g. Vail & Hardenbol 1979) it is clear 
thai posi Middle Miocene faulting is far more 
significant in the evolution of the cast Mt Lofty 
Ranges than suggested by Twidale & Bourne (1975). 
furthermore, the clear exposure of a reverse fault 
on the eastern margin of the Mt Lpfty. Ranges 
supports the notion of a compressive component 
to the forces involved in the Tertiary uplift of the 
eastern Ranges 

5 Lindsay, .1. M, (1986). New record and reclame 
implications of a high level f.epi(fo<vctirwbcxfwR 
ItnieMone, Bremer Valley, casrein Mount Lofly Ranges. 
S.A. Dtrpt of Mines and t-itcrity rcpori 86/66 (mipuhl,). 


This paper is published with pet mission of Ihe 
Director-General of Mines and hnergy, South 



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of the Mounl Lofly Ranges. Trims. /?. Sov. S. Aust. 35. 

i 'AMrANA. B. (1955) The structure of the LaUcnt South 

Australian Ranges; the Mount Uifty-OIary Arc X Oeol. 

SW- Aust. 2. 47-61. 
rmt'R A N (1964) Ternary lorarrnuilcra Irom 

Gippsland, Victoria, and their stratisrapfucal 

significance. Mem peoi. Sitrv. Vict. 23. 
t-iNNKR. (' (iy*0) The major structural uud 

physiographic lealuic* of South Australia. Trans. R. 

Sov. .V. Aust. 54, 1-36. 
Tikman, J. ft. (l%7) Stratigraphy of Late Cauwoic 

deposlrs in South Australia, /hid. 91, 165-17$. 
Ul apssvfr. M. K (1953) Some problems of TctUavy 

geology in southern Australia. Jour, Proc. /?. Soc, 

MS.W. 87, 31-45. 
I inosav. ,l, M. (1W5) Aspects of South Australian 

Tertiary Ibraniinifcral biosnatigraphy, wiih emphasis on 

studies of Massilina and Suhbotina. Spec: fHthi S. 
\ust Dept Mines c«dl £"e?g}\ 5. 187-232. 

Limwrook. N. H. (1961) Stratigraphy or the Murray Ba>hi 
in Souih Australia Bull. geut, Surv, S, Aust. 36, 

May, k, I, & Dourman, R. P; (1984) CoaslaJ 
iHndsHimpinjt in Pleistocene sediments at Sellieks fleaeh, 
■South Australia. Trons H Soc. S. Aust. 108, 85->M 

Srbkscl R. C. (1945) Sotne aspects of the geonjofnbolQfiy 
of portion of the Mount Lofty Ranges, /hid. 6«f, 

Twumu, C. R. & KooKNfc, J. A. (lsm> 
tieomorphologieal evolution of part of the extern 
Mount lofly Ranges, South Australia. Ibid. 99, 

Vaii, P. R. & HARorMrtni . J. (1979) Sea-level changes 
during rhc Teriiaty. Octanus. 22, 71-79. 

W'adi . M (1964) Application of the lineage eonccpl to 
bioslratisraphic zoning teased on ptanktonic 
foivurtinit'eia. Mii-ropuitfonrohxy III, 27^-290. 

Wiilman. P A CjRXrvHAroH, S. A. (J9R8) Hinders 
Mount bjfty Ranges. South Australia, their uplifl, 
erosion and relaiinnshin to crusial structure trans. H 
Soc. S. Aust. 112. 11-19. 



byR. C. Ainslie, Z). A. Johnston & E. W. Offler* 


The fauna of the intertidal mudflats of northern Spencer Gulf was monitored between 1982 and 
1986. Two main habitats were identified, the mid intertidal zone, and the seagrass fringing low 
intertidal zone. These habitats supported quantitatively differing faunas. 

There was no evidence of the species impoverishment reported elsewhere for the epizoic fauna of 
this region. Comparison of the characteristics of the fauna with that of another negative estuary, the 
Port River in Gulf St Vincent, has provided the basis of a monitoring programme to assess potential 
environmental stress imposed by power station development in northern Spencer Gulf. 
KEY WORDS: Intertidal habitats, benthic fauna, seagrass, Spencer Gulf. 



by R. C Ajnslie. D A. Johnston & E. W. OHi-.R* 


Aivtur, RjC.v Johnston. D.A. & Offler, E. W. (1989) Intertidal communities ol northern Spencer 
CutT, South Australia. Trans, R. Soc. S. Aust. 113( ), 69-8?, 31 May, I9S9. 

The fauna of the iiuertidal mudflats of northern Spencer Gulf was moniiored between 1962 and 19HG. 
TUo main hahitats were identified, the mid intertidnt zone, and the seagrass hiiujing low imetridal 7.onc. 
Thcse habitats supported quaiituaiivcly differing faunas. 

Trier.* Was no evidence of the species impoverishment reported elsewhere for the epi/oic fauna of this 
region Comparison of the characteristics of the trwna with that of another negative estuary, the Porr Riva 
in Cjulf St Vincent, has provided the basis o» a monitoring programme to assess potential environmental 
stress imposed by power station development in northern Spencer Gulf. 

Kr-v Words: Imertida! habitat^ bent bit. fauna, seagrass, Spencer Gulf. 


Northern Spencer Gull' can he defined as that 
ponton of Spencer Gulf, South Australia, north of 
the line from Ward Spit to Lowly Point, i.e. north 
of 33°S (Stcfanson 1977). It is a unique, narrow 
body or water, projecting inland some 200km from 
(he Southern Ocean coast (Fig. I), 

The waters of northern Spencer Gulf arc 
characterised by high summer salinities, up to 48 
(Nunes & Lennon 1986) and high summer 
temperatures with wide seasonal fluctuations, 
I1-25°C (Johnson 198P). Due to high evaporation, 
and low fresh water inflow to (lie norlhern Gulf, 
there is a longitudinal salinity gradient of about four 
in 20km. The region has been described as a 
negative estuary, 

A limited number of studies has been carried out 
on the marine communities ol* the "middle Gulf" 
(SEA I98I 2 ; Ward & Young 1982, 83) and concur- 

Johnson. J.fc. (1981) Hydrological Data for Upper 

Spencer Gulf 1975-1978. Fisheries kes. Paper No. 3, 

1-30, (Depi. l-'isheries, Adelaide), unpubl. 

Social and Ecological Assessment (19KI) Dmll 

Environmental Impact Statement for port and terminal 

facilities at Stony Point, South Australia. Prepared for 

SANTOS, unpubl. 

fCinhill (1987) Upper Spencer Cmlf Intertill Survey. 

Final Report, May 19H7. unpubl. 

THectridty Trust of South Australia (1977) Noithcrn 

Power Station Environmental Impact Statement. July 

(977, unpubl. 

Elecnicity Trust of South Australia (19*S5) Northern 

Power St jtion Environmental impact Statement. August 

1985. Prepared by Kinhill Stearns, unpubl. 

Environmental Sciences and Engineering Branch, 
Electricity Trust of South Australia^ P.O. Box n T 
Eastwood S. Aust. 5061 

rently with this study, Kinhill (1987)° have examined 
inte* tidal corn muni lies m Ward Spit and Lowly 
Point- Shepherd (1983) provides I he only published 
description of benthic communities of the narrow 
reaches of the northern Gulf. Shepherd found a 
marked reduction in species number in the epizoic 
communities on Pinna hicolor in northern Spencer 
Gulf compared to that of the same habitat in Gulf 
St Vincent, raising the possibility of a general 
species impoverishment fa the hyper-saline waters 
of northern Spencer Gulf. He emphasised the need 
for further detailed study of other "component 
parts" of the biological system to determine \\\ 
capacity lo accept additional stresses of industrial 
wastes and discharges. 

Since the mid 1950's Play ford Power Station has 
been operating on the eastern shore of the northern 
Gulf, discharging warmed cooling water to the Gulf. 
The discharge from Play ford Power Svtanon. 
operating 31 it's maximum nominal capacity of 
330MW, resulted in surface water temperatures 
about 6°C above ambient neai the power station, 
with only occasional incursions of warmed watei 
(about 3°C above ambient) to nearby intertidal 
zones (ETSA 1977 4 ). In 1977 and 1985 [fie 
Electricity Trust of South Australia issued 
Environmental Impact Statements fonhree further 
power station unils, with a total generating capacity 
of 750MW, to be built in this area. This 
development tonus the Not (hern Power Station. 
Thermal discharges were predicted to more Ihan 
double <FTSA IM77 4 , IW5*). 

The effects of warm water discharge from power 
stations on benthic fauna have been the subject ol 
a number of reviews (Coutant & Talmage 1975; 
Talmage & Coutant 1980; Craven et at 1983; 
Langford 1983). Effects vary from site to site as a 


R. C. AhNSLJE, D. A. JOHNSTON & t, W- Ol I I rk 

function of the climniic, hydrological and biological 
features (Crema & Pagliai 1980). Greatest effects 
are observed near the outfalls -at the hottest times 
of the year (Lungfurd 1983) and include changes 
in the biological characterisi ic* of the bent hie 
ujmrnuniiic.5 within the zone of influence of the 
warm water plume (Robinson et at 1983; Robinson 
1983; Thomas et af 1986). In particular. Hamper 
& Spencer (1984) and Thomas ef a! (1986) have 
described [he decline in populations of particular 
bivalve mollusc species and the establishment of 
dense populations o( opportunistic worm species 
as characteristic of thermal disturbance in soft 

The present study began m 1982, three years prior 
in Ihe commencement of operation of the new 
power station. Intermittent operation of the first 
250.MW unit commenced in the summer of 1985, 
with MXJMW operating by summer 1986. The aim 
ol the study w;^ to investigate spatial and LempoTal 
pal terns of 'he benthie communities in the negative 
estuary of northern Spencer Gull, it was anticipated 
rhat derailed descriptions of these communities 
would identify whether or not there were any 
biological impacts from the existing power Matron 
development (and early operation of the new 
development.), and establish baseline data for long 
term assessment of the effects of increased thermal 
discharge, This study concentrated on the intcrtidal 
and shallow sublittoral fringe, as the Only other 
published South Australian study on the impacts 
uTehcrtnal discharge on the marine environment 
had described well defined patterns of change in 
ihecnmmuuiiiesof this /one (Thomas H ol 1986). 

The influence on communities ofscduncni type, 
degree ol em erven* and presence of scagrnss arc 
investigated in this study. 

Station foevtions. elevations, and sediments. 

Near Port Augusta the narrow channel basin of 
northern Spencer (julf is bordered by shallow 
sublittoral seaumss meadows and extensive intcrtidal 
mudflats backed by stands of the jf«$| mangrove,. 
Avicontriis marina var resintjvra (Fig. I). 

Seven transects were established along the eastern 
shore of the gulf, both adjacent lo and remote from, 
existing and proposed power station developments 
(Fin. I). Three stations were established at di I fcn.ui 
inlcrhdal /ones nn each trained, Kach station had 
an urea of about 25m ? . On each transect, station 
I was located 30m seaward from the mangrove 
fringe, station s was located m the seaerass verge* 
(emergent at extreme low tide) and station 2 was 
located midway between I and 3. The locations and 
elevations nf each station w^ere confirmed relative 
to an established Port Augusta Powci Station 
datum, us!ii>* an fcDIVf theodolite. 

Sediment samples were collected from each 
.station on all transects and analysed for grain sivse 
according to the Wentwurih scale (Folk 1968) with 
minor modulations (Thomas et at 1986). Graphic 
mean*, Mz> (Folk L968) were deicrmincd for the 
samples. The Bray-Curtis classification technique 
was used to determine homogeneity groups based 
on the percentage distribution ol' particular 
sediment size classes in each sample (Miedeke & 
Stephenson 1977; Thomas et uf 1986). 

Water and air lempuruture. 

For the study period 1982-I9S6, a continuous 
record of Gulf water temperature was made at the 
cooling water intakes ol Play ford Power Station. 
A continuous three hourly record of ambient ait 
temperature was made ar iheSA Weather Bureau's 
Port Augusta weather station No. 019066. 


Each station was sampled using a hand operat- 
ed coret collecting a sediment sample of 16(10 cm-, 
surface area 80 ejnP, to a depth of 20 cm. The sam- 
ples were taken durin.e daylight hours. 

The pattern of species accumulation with repeat- 
ed random corine. within a station area was deter- 
mined. By subsamplc 2(1, about 90°ft of the spe- 
cies recorded in 40 sabsamples had been found, and 
those species which occurred after subsample 20 
uompr ised an eUrenrely small number of the tola! 
number of individuals sampled. The decision was 
made to ptocccd using 20 subsamples per station, 
exceeding the species-area requirements proposed 
by Cain & CastTO (1959) and meeting the stringent 
piopusals Of Weinberg { 1 97-8). Each station was 
sampled twice yearly (winter and summer) from 
June 19S2 to June 1986. Samples were treated as 
described in Thomas et af < 1986). Faunal relation- 
ships (between stations, transects, tidal /.ones, and 
sampling times) were interpreted using Bray-Oirtis 
classification utter root-root transformation of the 
data, and dendrograms were cojLstructcd with 
group average sorting (Swart/ 1978; Field et af 

Species constancy, abundance rank s-eoies, and 
average rank scenes were assigned as in Hailstone 
(1976) for 160 species totalling 34,3 J 2 individuals. 

Species associations were determined using Bray 
Curtis classification analysis on non-transformed 
data, considering the 40 top ranked species for the 
period 19S2-1986. 


Station elevations and sediments 
Tidal fluctuations relaiive to station tones result 

in dilferent periods of emergence for stations I, 2 
and 3 on all transects. Considering February 1 9S4, 



■50U1M '1e t 

500m | 

Fig. 1. Sampling station locations. Northern Spencer Gulf, South Australia. Inshore (rinsing seaarass species aUo 



l be station I zone is emergent for over 200 
hours/month, station 2 for about 80 hours/month, 
and station 3 for about 5 hours/ month, or less than 
1% of the time, In winter even shorter periods of 
emergence occur at the low intcrtidal zone. Local 
meteorological conditions, such as wind velocity, 
also affect the degree of exposure on low spring 
tides. Thus stations 1 and 2 are clearly mid inierti 
daJ stations while station 3 is in the low imcnidal 

The mid intertidal stations ate devoid of scagrass 
cover. The scagrass species at stations 3 include 
both Posidaniu australts and Povdonia sinuosa, 
without clear eul zonal ion in the shallow water. 
Some Amphibolus antanika is present at transect 


The hierarchical classification oi sediments from 
all stations gives two major groups o\ siaiion.%, 
pumps (a) and (b) Fig. 2. The sediments of stations 
forming group (a) have M/ values which generally 
range from medium sand (J. 5<z>) to coaTvc sand 
<0.25d). The exceptions are station U which is clas- 
sified as fine sand (M2 = 2.5'M, and the subgroup 
formed by sediments of stations 6. 1 and 6.2. these 
last two stations have M/. values between and 
-.25^ (ie. at the lower end of" the very cnatsc - sand 

The second major group of sediments (b) have 
Mz values ranging from very coarse sand (-.25<y) 
ihiough to granules (-1.7<£), with stations 7.1 and 
7.2 having the coarsest sediments; recorded in the 
survey. Although sediments of station 7,3 have an 
M/ value of l,25<6 (medium sand I, this station 
forms its own subgroup on the dendrogram because 
of 1 he restricted distribution in gram size around 
the median, unlike all other samples which had 
mure 01 less even ranges of grain sizes. 

Air and water temperatures 

Air and water temperatures 1982, 1964 and 1985 
show that seasonal water temperatures follow air 
temperatures. Amhieni average monthly wHier tem- 
peralures range from U-I2,5°C in winlei to 
24-25 °C in summer (Fig. $). 

I here is- no indication of change In the overall 
ambient temperature from 1982-1985, as a result 
of intermineni cooling water discharge from the 
existing power station development. 


A total of lrX> species were recorded ; polyehaetes 
were ihe most numerous with 52 species, includ- 
ing several new records for S, Ausa. (Tabic I). Mol- 
luscs with 50 species and crustaceans, 40 species. 
were nest in abundance, with 1$ species from var 
ious other ta\a < Table t). 


* 90- 

1 ^ 







fig. 2. Dendrogram ni northern Spencer Gull' nations liy 
sediment grades. 

Fig. 4 presents the station faunal homogeneity 
dendrograms for the first two seasonal surveys 
(winter 1982, summer 1983), and for the last two 
surveys (summer and winter 1986). In all cases, at 
about the 25-30% level the homogeneity dendro- 
grams consistently show two major collection 
clusters with high infra-group homogeneity (Fig. 
4 (1-4), Groups A and B). Generally the faunas 
of stations 1 and 2 on all transects comprise Group 
Aon each dendrogram, and siation 3 faunas form 
Group B. In two of the dendrograms presented 
(winter 1982 and summer 1986) there are exceptions 
to this generalisation but there is no seasonal ot 
spatial patict n to the exceptions, and the major 
trend persists. There is no evidence of consistent 
subgrouping within the major observed associations 
(Group A and Group 0, Fig. 4). This tendency for 
launas of mtd tntcnitial stations 1 and 2 10 group 
more closely togetliei than to those of station 3 has 
been consistent throughout the lour ycai study peri- 
od and is also reflected in the the lidal zone analy- 
sis, (Fig. 5) for each survey 1982-1986. 

The species homogeneity dendrogram deter 
mined using ihe 40 most common species is present- 
ed in Fig. 6. Accepting the arbitrary cut-off level 
of 25% similarity, this dendrogram shows two 
major groups of species (Al and Bl, Fig. 6). 
Although all but two of the forty most common 
species occur over all stations, those which form 
Group A I always contribute a grealei proportion 
of the total number of individuals in the samples 
of stations I and 2 and those of Ciioup Bl arc al- 
ways more abundant in the station 3 faunas, 

Together these analyses (Figs. 4, 5 & 6) indicate. 
'hat the two habitats represented by the mid bi:cr 
tidal region (stations I and 2) and the low inteni- 


Table 1. Species recorded throughout the stud y. New records for South Australia are indicated by an asterisk. 

Exclusively Exclusively 

mid intertidal low intertidal 


Qiganereis edmondsi (Hartman) 

Neanthes vaalii Kinberg 

Neanthes cricognatha (Ehlers) 

Ceratonereis transversa Hutchings & Turvey + 

Ceratonereis miraoiiis Kinberg 

Perinereis nuntia (Grube) 

Perinereis amblyodonta (Schmarda) + 

Perinereis sp 

Nereis triangularis Hutchings & Turvey 

Nereis sp + 

Nematonereis unicornis (Grube)* + 

Marphysa sp* 

Arabellidae 1* + 

Arabellidae 2* + 

Lysidice sp 

Eunice sp + 

Schistomeringos sp 

I .umbrinereis sp 

Nephtys austraiiensis Fauchald 

Gtycera americana Leidy 

Phyio sp* + 

Scoloplos cylindrifer Ehlers 

Leitoscoloplos normalis Day* 

Leitoscoloplos sp 

Naineris grubei australis Hartman 

Phyllodocidae 1* + 

Phyllodocidae 2* + 

Phyllodocidae 3* + 

Cirriformia Jiligera (Grube) 

Cirratulidae sp 

Boccardia chilensis Blake & Kudenov 

Harmothoe sp 1 A + 

Harmothoe sp 2 A + 

Harmothoe sp 3 A + 

Lysilla apheles Hutchings* 

Liomia medusa (Savigny)* 

Eupolymnia nebulosa (Montague) 

Streblosoma sp* 

Terebellinae 1 

Terebellinae 2 + 

Terebellinae 3 + 

Terebellinae 4 + 

Polycirrinae* + 

A - species seperated on basis of anterior elytrae. 
Harantolla lepte Hutchings 

Notomastus sp + 


Pherusa sp + 

Armandia sp 

Gaieolaria caespitosa (Savigny) + 

Scalibregma sp* _$. 

Magelona sp* 


Nemertean spp + 


Golfingia margaritacea adelaidensis Edmonds 



mid intertidal 

low intertidal 


Teilina deltoidafis Lamarck 

Laternula recta (Reeve) 

Kateiysia peronti Lamarck 

Katelysia scalarina (Lamarck) 

Brachidontes erosus (Lamarck) 

Xenostrobus inconstans (Lamarck) 

Ostrea angasi Sowerby 

Pinna bicolor Gmelin 

Malleus meridianus Cotton 

Anomia trigonopsis Hutton 

Circe scripta Cotton 

Electroma georgiana (Quoy & Gaimard) 

Trichomya hirsuta Lamarck 

Chiamys bifrons Lamarck 

Musculus paulucciae Crosse 

Corbula jlindersi Cotton 



Salinator fragilis (Lamarck) 

Nassarius burchardi (Philippa) 

Nassarius pauperatus (Lamarck) 

Monodonta constricta Lamarck 

Bembicium melanostomum (Gmelin) 

BatiUaria estuarina (Tate) 

Batillaria diemenensis (Quoy & Gaimard) 

Bedeva paivae (Crosse) 

ComineUa erburnea (Reeve) 

Cominella sp 

Austroiiotia densilineata (Tate) 

Diala lauta A. Adams 

Quibutla tenuissima Sowerby 

Lepsiella vinosa Lamarck 

Conus anemone Lamarck 

Cantharidus irisodontes (Quoy & Gaimard) 

Clanculus ptebejus (Philippi) 

Clancuius weedingi (Cotton) 

Phasianetla australis (Gmelin) 

Hydrobia sp 

Microcolus sp 

Sophismalepas nigrita (Sowerby) 

Pterynotus triformis (Reeve) 



Notomella sp 

Elegidion occiduus CoUun 

Asteracmea sp 1 

Asteracmea sp 2 

Stomatella auricula (Lamarck) 


Ischnochiton variegatus Adams & Angus 
Ischnochiton contractus Reeve 
J-feterozona sp 


Hapalochlaerw maculosa Quoy & Gaimard 


Exclusively Exclusively 
mid intertidal low intertidal 



Penaeus latisuleatus (Kishinonye) 

Processa gracilis (Baker) 

Leander serenus (Heller) 

Leander intermedins (Stimpson) + 

Paguristes frontalis (M. Edwards) 

Paguristes brevirostris (Baker) . 

Caliianassa aequimanu (Baker) + 

Callianassa ceramica (Fulton & Grant) 

Crangon socialis (Heller) 

Pilumnus fissifrons (Stimpson) 

Philyra laevis (Bell) 

Cryptocnemus vincentianus Hale . 

Polyonyx transversus (Haswell) . 

Gomeza bicornis (Gray) , 

Halicarcinus ovatus (Stimpson) 

Litocheira bispinosa (Kinahan) I 

Portunus pelagicus (Linnaeus) + 

Helograpsus haswellianus (Whitelegge) 

Ceratoplax punctata (Baker) + 


Maera mastersii (Haswell) + 

Birubius panamunus Barnard & Drumrnond 

Etasmopus bampo Barnard 

Allorchestes compressa (Dana) 

Ceradocus serratus (Bate) + 

Cymadusa sp 

Gammaridae 1 

Gammaridae 2 

Gammaridae 3 + 


Cymodoce longicaudata (Baker) + 

Zuzara venosa (Stebbing) 4. 

Exosphaeroma sp + 

Cirolana woodjonesi (Hale) i 


Anthuridae 1 

Anthuridae 2 


Apsuedes australis (Haswell) 
Paratanais ignotus (Chilton) 


Halanus amphitrtte Darwin 
Elminius modestus Darwin 


Anthothoe albocincta (Stuckey) 

Cricophorus nutrix (Stuckey) * 


Nesogobius hinsbyi (McCulloch & Ogilby) 
Gymnapistes marmoratus (Cuvier & Valenciennes) 



mid intertidal 


tow intertidal 


Lvptasynapfa dufobrijera (Stimpson) 

Taemogyrus mebucki (Jos-luia) 

Arrthtypneustes sp 

Trochodvtu sp 

Ophiuroidea 1 

Ophiuroidea 2 




Ascidta sp 

dat fringe (stations 3) can be considered to have 
quantitatively differing faunas. 

Fig. 7 compares mean number of speeies/m 2 , 
and mean numbers of individuals/m 2 for these two 
habitats over the period 1982-1986. Significantly 
more species/m 2 are found in the low intertidal sta- 
iions. Reference to the species list for all surveys 
reveals that of the 160 species collected, 86 (54<7o) 
occur over the whole tidal range. This 55^o includes 
37 of the 40 most common species, Fifty-nine spe- 
cies, or 38% of the total* occur exclusively in the 
low intertidal zone, and only 8% occur exclusively 
in the mid Lnteriidai /one occupied by stations 1 
and 2 (Table 1). In contrast to species abundance, 
the communities of mid intertidal stations 1 and 
2 consistently have significantly higher mean num- 
bers of individuals (Fig. 7) than those of the 
scagrass fringing stations 3. Nine of the 10 most 
common species are more abundant at stations 1 
and 2 (Table 2, Fig. 6); conversely, of the 61 spe- 
cies exclusive to Ihe station 3 community, only two 
appear in the "top 40" list (Table 2). Nevertheless 
the less common, but exclusively station 3 species, 
contribute to the consistent separation on the sta- 
tion and tidal zone faunal homogeneity dendro- 
grams (Figs. 4 and 5). 

big, 8 shows (he intersurvey time dendrograms 
comparing whole survey collections for each of the 
two communities for the period 1982-1986. 


CornmwiUy-hubitat relationships of ihe mid inter- 
tidal and fow intcrlidul zones. 
In many studies sediment characteristics have 
been shown to be a significant factor in determin- 
ing bcuthic community patterns (Gray 1974; 
Saertger el al 1982; Jones et al 1986). In intertidal 

regions particularly, there is the potential for wave 
action to grade sediments, resulting in coarser par- 
ticles on the upper shore line. Such a graded series 
of sediments is potentially reflected in the nature 
ofthebenthic communities (Gage 1974; Robinson 
1985). On the low energy shore of northern Spencer 
Gulf there is no sediment gradation, with random 
groupings of station sediments irrespective of the 
zones in which the stations occur. There is a high 
overall similarity in the sediments, the two main 
groups discerned (medium to coarse sand, and 
coarse sand to granules) showing about 70 < Vo 
similarity on the homogeneity dendrogram (Fig. 2). 
In common with other studies of the South Aus- 
tralian Gulfs where a limited range of sediments 
is found (Kinhill 1987 3 ; Thomas et ul 1986) there 
is no evidence that this factor influences benthic 
community distributions in the northern Spencer 

The mid intertidal zone is characterised by spe- 
cies which, although more common in this /one, 
occur Over the whole tidal range (Tabic 1). The 
most abundant species is the mussel, Bruchidontes 
erosus? which forms dense mats in this zone. Such 
mussel mats are characteristic of many low energy 
sand and mud Hat substrata in S, Aust. (Womerv- 
ley & Thomas 1976; Thomas et al 1986). Other 
bivalves, although less abundant than the mussels, 
are also characteristic of the mid intertidal zone. 
These include Tellina deltokialis, a species belong- 
ing to a genus known to have a behavioural 
response to light direction which contributes to its 
zone maintenance (Newell 1979), Katetysia scala- 
rina, and Laternula recta. These bivalves contrib- 
ute to the abundance of many other species as they 
provide the substratum for the settlement of irur 
barnacles Elminius modestus and Bahmus ampfa- 











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iUH JUL *uC u* [>;■ K-rv c«' 

Fig. 3. Monthly averages of daily maximum air 
temperatures recorded at SA Weather Bureau Station 
No OI90to, Playford Power Station, Pt Augusta {Tig 
I). Monthly average ambient Gulf water temperature 
derived from continuous hourly records at the 
circulating water intakes, Playford Power SUition. 

trite, and the sea anemone Anihoihoe afbocincta, 
There also appears to be a close relationship be- 
tween the large bivalve species and the omnivorous 
worm species Nephtys austral iensis, Neanthes 
vaalii, Olganereis edmundsi, and Nereis trian- 
gularis. The last species seems generally to be as- 
sociated with mussel beds (Hutchings & Murray 
1984). Several species of scavenging and car- 
nivorous snails (Nassarius pauperatus, Bedeva pai- 

vae> Batillaria estuarina, and Batillaria diemtnen- 
sis) also occur in the community dominated by the 
mussel beds, as docs the scavenging crab Philyra 
iaevis, characteristic of the mid intertidal zone on 
low energy coastlines of S. Aust. (Womersley & 
Thomas 1976). The herbivorous top shell, 
Monodonta constricw is common among the dense 
mussel beds which provide a substrate for algaJ 
growth, particularly Enteromorpha spp. 

In the extreme weather conditions of the north- 
ern Spcncei Gulf it is clear that a number of ubiq- 
uitous intertidal species are restricted to the low in- 
tertidal zone. These include the polychaetes 
Nemmonereis unicornis, Ceratonercis transversa, 
Baranfo/la lepte f Cirriformia fili%era and 
Leltoscoloplos norrnalis % for which a range of 
habitats has been documented (Hutching* & Mm- 
ray 1984) and the gastropods Nassarius burclwrdi, 
and Austrolioiia densilineata> also found elsewhere 
over broad tidal ranges (Thomas et al 1986), Con- 
versely, there are a number of species which are 
generally characteristic of the subtidal zone. These 
include such .species as the hammer oyster Malleus 
meridianus, the scallop Chlamys bij'rons, the blue 
swimmer crab Portunus petagicus. and several 
other species primarily, or only, found in associa- 
tion with seagrass (Electrotna f>eorgiana (Ludbiook 
1984), Gol/ingia margaritacea adelaidensis [Ed- 
monds 1980) and Amblypneustcs sp. (Shepherd A 
Sprigg 1976)). The major environmental factor 
which appears to separate the habitats described 
in this study is the "degree of exposure". This is 
a function not only of the length of the periods of 
emersion (in a region of high summer air tempera- 
tures) but also the protection from environmental 
extremes provided by the presence of seagrass in 
the low inLertidal zone. 

Comparisons with other areas. 

Are the intertidal communities of northern 
Spencer Gulf impoverished as evidence may sug- 
gest for cpizoic fauna on Pintw tricolor? There are 
inherent difficulties in comparing communities of 
various marine and estuarine areas. Generalisations 
from one area to another are often of limited value 
beyond the broad observation that marine domi- 
nated sites generally have more species than estua- 
rine sites (Collett et al 1984). Accepting this limi- 
tation it can nevertheless be seen that the total num- 
ber of species found in the present study (160) is 
within the range of species number recorded in 
several studies of eastern Australian "estuarinc" 
regions (Rainer & Fitzhardinge 1981; Jones ei al 
1986), and is comparable to the number recorded 
in a tecent study of the intertidal mudflat fauna 
of the Port River estuary. Gulf St Vincent (Tho- 
mas et al 1986). About one and a half times as 








30- , 

1 1 

> "°- 

1 rii n 
1 n 

s 50- p- 1 — ] 

It n M] 

" 70- (| 

dti r 1 ] 




5 50 


£ 60 


5 70- 












A B 

Fig. 4. Station faunal homogeneity dendrograms. (1) winter 19X2, (2) summer 1 ( JX3, (3) summer 19W\ (4) winter 1986. 

many species are recorded in the present study as 
were recorded by Kinhill (1987)3 for intcrtidal mud- 
Hats about SO km further south in Spencer Gulf 
where salinities are four to five lower (Nunes & 
l£nnon 1986). Kinhiil (1987)^ did not sample at the 
seagrass fringe (D Evans pers. comm). Disregarding 
the 38°Fu of the total number of species which arc 
exclusive to the seagrass Cringe in the present study 
leaves a species number very similar to that found 
by Kinhill (1987) 3 . Therefore this study provides no 
evidence of species impoverishment in the imertidal 
zone, related to high salinity or other environmental 
factors characteristic of the area. 

Natural variations in the interridal communities of 
northern Spencer Gulf 

For a baseline study to be of use in post develop- 
ment monitoring an understanding of natural var- 
iations in the species matrix is necessary, The faunal 
homogeneity dendrograms for both stations and 
tidal zones demonstrate the consistent relationship 
between the mid intertidaJ and low intertidal com- 
munities over several seasons. Between 1982 and 
1986 the common core species characterising these 
two habitats remained constant (Tabic 2). Perhaps 
surprisingly no seasonal pattern emerged from the 
intersurvey comparisons. The overriding pattern 


which showed for both habitats was the trend for ovcraJi similarity in survey faunas over time this 
sequential subgrouping ol* surveys, with survey fau- seems the most likely explanation here. The slightly 
nas within each of the two habitats showing high lower intersurvcy similarities of the low intertidal 
similarity, grouping at the 60°7u-7G^o similarity lev- communities, compared to those of the mid inter- 
el in each case. Sequential grouping of survey fau- tidal communities, is a consequence of the fact that 
nas does not appear to be unusual. Saengcr et al the former are characterised by more rare or occa- 
(1980) interpreted such a pattern as demonstrating sional species, 
progressive recolonisation following environmen- 
tal disturbance. Buchanan et al (1974, 1978), Monitoring for changes associated with environ- 
however, observed similar sequential subgrouping mental disturbance 

of survey faunas from undisturbed marine environ- Changes in estuarine faunas in response to or- 

ments and interpreted this pattern as indicating a ganic or thermal disturbance have been well 

more or less continual change in the presence and documented (Gray 1976; Parker 1980; Bamber & 

abundance of occasional or rare species while the Spencer 1984; Thomas et al 1986). In particular, 

core species remain relatively constant. Given the in the negative estuary of the Port River, Gulf St 

TABLb 2. Common species of the intertidal fauna of Northern Spencer Gulf, determined by reference to constancy, 
and abundance rank scores (Hailswne 1976}, Note 1 hat the top 40 species ranked in this table includes all 21 species 
which would he selected on Field et a/'s (1982) arbitrary cutoff for dominant species, namely species which contribute 
4^o or more to the total population or any one survey, 


{Refer Fig 6) 

Olgartereis edmondsi 
Nephtyx australtensis 
Braehidomes erosus 
Tel/ina delroidalis 
Nassarius pauperutiix 
Paguristes frontalis 
Baullaria diemenensis 
Katelysia scalurina 
Elminius modes tus 
Scnloplas cyiindrifer 
Neanthes vaalu 
Monodonta constricta 
Callianassa ceramicu 
Salwator fragitis 
Cirriformia filigera 
Batanus amphitrite 
Baraniolla lepte 
Apsuedes auslratis 
Nemeriean spp 
Nereis triangularis 
Nassarias burchardi 
Marphysa sp 
Terebellinae 1 
Lysifta apheles 
Crangon sociaiis 
Phtlyra taevis 
Laternula recta 
Asteracmea sp 1 
Batillaria estuarina 
Bedevu paivae 
A mhothoe albocincta 
Eupoiymnia nebutosa 
Strebiosoma sp 
Katelysia perortii 
Gotjingia m adefaidensk 
Elect roma georgiana 
Austroliotia densilineata 
Schisiomeringos sp 
Leitoscotoplos normalis 

Most common species in the low intertidal zone Group Bl I-ic. 6. 








°6 * 
















fl H 









CI 8 






























































































































10U2 1083 1863 

10B« 1B84 1985 1985 i960 10 86 




" ;o 





e^ 00 

j^J 3 123 123 123 123 123 123 123 123 



Fig. 5. Faunal homogeneity dendrogram* with respect to tidal zones for all surveys, 1982-1986. 

W1 ft* W3 B7 •! ■> W* C7 I1fl W31 tn X* BJ WIO C4 wit ci« cm wat mo 
. ft13 W11 M WJ CI W« 111 914 C» •!« Wit C31 W7 W30 WM ft* ftSS W33 W20 WIS 

v / V i- - ' 




Fig. 6. Species homogeneity dendrogram determined on non transformed data for the 40 most common species for 
the period 1982-1986 as determined by average rank scores assigned as in Hailstone (1976). 









1 | 










1 1 



















1 A2 


Fig. 7. Comparison of numbers of specics/m 2 and 
numbers of individuals/m 2 for mid intertidal stations 
I and 2, and for low intertidal stations 3%. 

Vincent, Thomas et al (1986) describe responses of 
intertidal communities to progressive increase of 
thermal discharge from Torrens Island Power 
Station. These include suppression or elimination 
of populations of bivalve mollusc and worm species 
characteristic of undisturbed regions, and increases 
in populations of opportunistic worm species. Of 
the 120 species recorded from the Port River estuary 
about 50 species (40%) also contribute to the 160 
species of the present study. These include several 
bivalve and worm species which had a negative 
response to thermal discharges, and which played 
a significant part in defining thermal perturbation 
in the Port River, namely the bivaJves Tetlina del- 
toidalis, Kateiysia scalarina and A', perortii, and the 
worms Nephtys australiensis, Neanlhes vaulii and 
Scolopfos cytindrifer. All of these species are 
common in the communities of the mudflats of 
northern Spencer Gulf, particularly in the mid 
intertidal zone (Table 2), The presence of these 
common indicator species provides the basis of a 
monitoring programme to detect potential thermal 
perturbation in northern Spencer Gulf. Any 
changes in abundance of these species in areas near 
thermal discharges will be reflected in recognisa- 
ble changes to the stable faunal homogeneity rela- 



> 70 - 






12345 6789 







JH 70- 




# 90 


Fig. 8. Intersurvey faunal homogeneity dendrograms, (a) 
staiions 1 and 2 r (h) stations 3. Survey number I -winter 
1982, 2 - summer 1983, 3 - winter 1983, 4 - summer 
1984, 5 - winter 1984, 6 - summer 1985, 7 - winter 1985, 
K - summer 1986, 9 - winter 1986. 

tionships described in this study over the period 
1982-1986. Such changes in faunal homogeneity 
dendrogram patterns have been clearly demonstrat- 
ed by Thomas el al (1986) in studies of changes in 
the fauna of the Port River estuary with progres- 
sive increase in thermal discharge to that region. 
The cirratulid worm Cirrifomia fiiigera is common 
in the low intertidal zone (Tables 1-2). Although 
the life history of this particular species is not well 
known, other members Of this genus have been 
shown to have typical opportunistic characteristics, 
capable of rapid colonisation of disturbed regions 


K. I ., ». A. lOHKSION A f. W. DjTLER 

(Gecnge 1964 a,b; Tbornas pi a/ l*>86). Jhe ar- 
chetypal oppoiiuiiisue worm species CaviteUa 
capitatu {Grrt$le& Grassier 1974; Gray 1976; TsM* 
Band 1987) has not been recorded hi the |>i«sent 
study. It is. however, present in intertidal commu- 
nities 50 km furthet south (Kinhill 19873), an d it 
is not unusual lor opportunistic species in disturbed 
regions to be immigrants from other fegions (Trio 
mas ft at \9H6). As with changes in the indicator 
species discussed above, any changes in populations 
ol opportunistic worm species- near the thermal dis- 
charge can be expected to noticeably alter the pat- 
ients or the faunal homogeneity dendrograms, 
waning of potentially significant environmental 
perturbation, lo WM no impact ol tltc occasion 
al incursions ot warmed water from Thomas Play- 

lord Power Station to the intertidal zone, and the 
initial discharge from the Nouhern Power Station 
(from summer 1985) can be discerned from com- 
munity relationships described in the present study. 


This work was funded and supported by ibe 
Electricity Trust Of South Australia as part of its 
ongoing programme of environmental research. 
We are grateful to those who idejitlilexi organisms 
especially Or R Hutching* (polychaetcs) and Mr W. 
Zeidler (amphipods) and to the South Australian 
Museum for access to their relerencc collections. 
Wc also thank Dr M, Geddes for his suggestions 
Mid comments, 


RAMIirfe, K. N. & SPHNtf*;£ h tH84) The benthos oV 

a constat powei station theenal Jisvhatuecatial. J. Mar. 

trial Ass. U.K. 64, 6Q9-42& 
MmihasavJ, B. & Wmwick K, M 11974) An estimate 

of bcmhie maetol'aunal pioduvuon in the offshore mud 

of the Northumberland coast. Ibid. 54, 197-222. 
Shmoi-k M„ ii Kinosion, P. F <l l )78) Sow cm 

of variability m rho henthic maeiolauna off tHti South 

Northumberland const, 1971-1976. Ibid. 191-209. 
C\in. Si A. &. CXm m G. M., ot- O. (1959) "Manual ol 

Vegetation Analysis" (Harper, New Y*mK) 
t'oLt rrr. L, G . Hv"rcitiNos. P. A., Gibbs. P. J. <k CM i »n\ 

A. ). (1984) A comparative study of the rnaerohenrhic 

fauna of Poiitdonta uimrahs seagrass meadows in New. 

South Wales. 40IW/C &Qt. 18, 111-134, 
C'OoiANt.G C, &Ta» MAi.h, S. S. (1975) Thermal effects. 

J. Hair* Holt. Control fttf. 47(6), 1657-1711. 
CcwtV, J. B . HWWEtSON.M. E. A T,U.MAGE,S. X 1*198.1) 

Thcnnal effects, fhid. 55(6), 787-800. 
i.Ut ma, R. £ I'Ai ;UM. A. M. B, (14X0) The structure ul' 

hem hie couiuitmitie* in an urea of iheunal discharge 

from u coastal power station, Mar, Poitur. Hull. II, 

h'DMOsms, S. .1. (1980) A revision of the jyucmatktt of 

Ausuaiian sipuneulans tSipuneula) Rcc. S. Aunt. Aftts. 

1811), I -74. 
I ii i i), J. (i., O ark K. R <fc Warwick. K. M. (1952) A 

pitteOe-ttl .sitHiegy tor atialvsinu muhnpeeics distribution 

patterns. Mar ficot. Prog, .Scr H, 17-52. 
lot*., K. I.. (IWH) "Pc-nolop.y ot Scriimcwaiy Rocks" 

(Hemphill, levv), 
r;,viu J. tl9?li Shallow-water /-onation el Sea Loch 

benthos and its relation to hydrojrraprnc and other fcarmv..v J. Mar, btof Ass. UK. 54, 223-249. 
GeorOP. J. U <J>>6'1»> The lite history ol the nrratulid 

worm CtrrJfonnia (etuaculatu em an intertidal mudflar. 

;*lrf 47-65. 

(1964b) Ou some eiiviroiunenia! factors affecting 

die distribution of Cirnfar/niu tentaculuta tKitvchacu) 

at Hi.nil5le th\a 44, 373-5HK 
Ora>s* r- ( 3 F & ORA^SLfc J. P <I97/1l Opportunistic life 

hiiturio <ind genetic WMcrm in marine henrhic 

polychuetcs. ./. Mar. Res. 12<2), 2M-2K4. 
U6av I. S. yi974) Animat-scdinicm fclationstnps, 

ih-*>utn>)>niphy nrtd \futitt? Biol. Atw. Rev. it 223-261. 
(1976) The fauna of the polliued Rivet t.ees 

t'jtfuary. Fstuar. t rtdflf. Mm*. V7. 1, 203-23), 

llAlLsroSE. T, & (1976) Ddimitalion ol stibtkial 
macrohentlio.s associations at the mouth of the Drisbcme 

Kuet 4«4r- 7 Milt fresh*: Ht's. 27, 217-38 
Hih MINOS H A. *& Mi kkav A. (1984) Taxonomy of 
polychaetes from the Hawkesbury Kiver and the 
:iOuthe^n estuaries of fsiew Sonrh Wales, Austi'HUa. /fcv. 
Atfil Muy 3fe, (suppl & T-118. 
.Ioms A. R., Wmson Ri ^shti, C. J.. & Mokkv, A. 

I iv.sfo Sj\uial iMttcrns in the macrnhcnihic communities 

of the Hawkeshury fcstuaiv. New South Wales, ^wr 
1 Mar. Wwlw, Jto. 37. 521-43. 
LavjH'Ko, T. E. 1 1983 1 "Tleciiicitv feneration and the 

hcoicf y of Natural Waters' fl tverpoot Unoorsity Wf$9, 

London I 
LoofiKt-H>K,N-H. (1084) "Quaternary Molluscs of South 

Australia" Department of Mines and tuergy South 

Australia Handbook No. 9 D.j. (Woolman Govt Piittler. 

Mtt orcM , J. <V & StrpiirNyrN, W. (1977) Lnvironmentnl 

Intpaei M'td baseline studies of the sof( bottom marine 

environment ut the vidnity of tKcuu sewer outfalls on 

the NSW central coast. Third Au\t. Conference of 

CVoMal and Ocean Kngineertng I57-Ifi2 (liisMutioii 

of Engineers. Austratin, Melbourne), 
Ntwitt.R. C. (1979) "Hiology o) Intertidal AniftwU". 

(3rd cd.> (haueishnrn, U.K.). 
N'iM\R. A. & Lr:\NwN.G. W. (19S6I Physical piopaty 

diwriOnnotH and seaMonai trends \n Spencer Gulf. South 

AnsrraJia: an inverse estuary. Aitfi. J. Mar. Frvihw Ret. 

37(1), >9-53. 
P\MKtt* ( I G. (1980) tiliats of pollution Upon |h<- 

benihoiof Dellasl Ukirii. .Mi/'; r\>ti t Bull, II, 80-KJ 
k.MNik S. &. Fii/iiAuoiMit,, R. C. (1981) BeiUhie 

coiiitnuiiiiie^ in an esiuaiv with periodic deoxvvennnon. 

AusL J. Wffft freshw. Rt>y 32. 227-43. 
Kt.finoON, K. I. M. (1985) Temporal changes in the 

bemlue tttaeiolauna ol Lake MaequHiK', a New Soull' 

Wales coanal Uieoon. Wethvtds 5(2), 78-90. 
Van Or ft Vu oi , J. T, & Goms. P. .1. (1983) A 

Mitvi.y ol tin' estutione benthic fauna of Homchush Hov, 

rarramatia kiver. N.S.W. fHM 3(2), SI-K4. 
S\r'.ci>.. K, STri'ttnNsoN, W, & Movimiv, .1. (1980) The 

evtuaiiiie iTia^obenthos of the Calliope River and 

AuekUuO Cieek. QtM.vnMi*u*1, Mc*» QKl &$**, 2W(M» 

, i &. (!yS2) Macrobemhos ofihp 

cooling wuter discharge eanal of the GtaOMOue Powei 



Station, Queensland. Aust. J. Mar. Freshw. Res. 33, 

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Thomas, I. M., Ainslie, R. C, Johnston, D. A.,, 
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Ward, J. J. & Young, R. C. (1982) Effects of sediments, 
trace metals and particle size on community structure 
and epibenthic seagrass fauna near a lead smelter, South 
Australia. Ibid 9, 137-146. 

, & (1983) The depauperation of epifauna 

on Pinna bicolor near a lead smelter, Spencer Gulf, 
South Australia. Environ. Poll. (Set, A) 30, 293-308. 

Weinberg. S. (1978) The minimal area problem in 
invertebrate communities of Mediterranean rocky 
substrata. Mar. Biol. 49, 33-40. 

WoMERSEhv, H. B. S. & Thomas. I. M. (1976) Intertidal 
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B. P. Webb (Eds), "Natural History of the Adelaide 
Region". (Royal Society of South Australia, Adelaide). 




BY W, KOSTE* & R. 7. SHIELf 


Diagnostic keys are given to the Australian representatives of the Rotifera: Monogononta in the 
families Euchlanidae (Manfrediutn, Diplois y Dipleuchlanis y Tripleuchlanis y Euchlanis) y Mytilinidae 
(Mytilina y Lophocharis) and Trichotriidae (Wolga y Macrochaetus y Trichotria). All species known 
from Australian inland waters are described and figured, as are some widely distributed taxa not yet 
recorded from the continent. Distribution data and ecological information also are given. 
KEY WORDS: Rotifera, Euchlanidae, Mytilinidae, Trichotriidae, Australia, taxonomic revision. 




by W. KbSTB* & R. J. Shieli 


Kostf, W. Sl Shii-.l, R. J. (1989) Rotifers from Australian Inland Waters. 111. Euchlanidae, Mytilinidae and 
Irichotriidae (Rotifera.-Monogononta). Trans, R. Soc. Aust. 113, 85 114, 31 May, 1989. 

L)iH gnostic keys are given to ihe Australian representative* nf the KoiilerfliMotioyononif* in ihe families 
Euchlanidae (Mar\f nullum. Diploic Diphuchlanis, Trtplcuchlamx, EuchhntsK Mytilinidae (Myltlintt, 
Lophochans) and Tnchotriidac {Wnlga, Macrochaetus, T/tckntrra/. All species known from Australian 
inland wuiets arc described and figured, as are some widely distributed ttxd not yet recorded from the 
continent. Distribution data and ecological information also are given. 

KCY Word';; Rotitera, Euchlanidae, Mytilinidae, TrK'hotHiilae. Australia, laKpnornic revision, 


The families of Rotifera;Monogononta 
considered in this pan of our review of the 
Australian species (sec Koslc & Shiel 1986b, I987h) 
commonly are encountered in the shallow littoral 
zones of billabongs or vegetated lake margins (cf. 
Koste 19X1, Tail et al. 1984, Koste & Sliicl 1986a, 
1987a, Koste et aL 1988). Some species, eg, 
Euchlanis dikttata and related forms, are known to 
become planktonic in eutrophic waters during 
cyanobacterM blooms (Ruttner-Kolisko 1974). This 
Species occurs seasonally in trie lower R. Murray, 
S. Aust. (Shiel et al. 1982). Otherwise, specks or 
the Euchlanidae (Manfredium, Diptois, 
Dfpteuchianis, Triplcuch/ani.s, EucbJctnis), 
Mytilinidae (Mytitina, Lophocharts) and 
Trichoti'iidae (Hxtlgu, Macrochueius, Triehotriu) 
generally are benthic or epiphytic in habit, and 
occur in open water of lakes, reservoirs or rivets 
only as incursion species during .seasonal flushing 

Most of the 35 taxa recorded m the three families 
were collected as such incidental incursions into the 
limnetic region of billabongs. It is likely that more 
taxa remain to be described, particularly endemic 
species, because our initial sampling program was 
directed al the planktonic community and the 
sampling bias has "under sampled*' Ihe preferred 
habitat of these littoral rou'fers. Only four endemic 
species are known in the three families, one 
mytilinid {Lophovharis curvata) and three 
triehotriids (Trkhoirkt pseudocurta and T. buehneri 
from Tasmania, and Macrochaetus danneefi, from 
ifc NT.). In contrast, about 25"7o of the taxa in the 
Epiphanidae and Brachionidae (Koste & Shiel 

• Rontgcnsir. 1, 2903 Bad, /wjwhenahn. Fed. Republic 

of Crermany 
fMurray- Darling Freshwater Research Centre, PjOi Bo* 

to, Alnury, N.S.W. 2640 

1987b), true limnetic species (and therefore more 
intensively sampled), appatemly are endemic. 

All known Australian representatives of the three 
families are described and toured, and diagnostic 
keys are provided to enable identification to species. 
Widely-distributed taxa not yet recorded from 
Australia, hut which are likely to be found here, are 
included in some genera. The format follows that 
of earlier papers; dichotomous keys arc followed 
by individual species" descriptions and known 
Australian records. Relative abundance from >20O0 
collections is indicated by: 'Yare" (fewer than ten 
records, "uncommon 1 * (10-30 localities) and 
"common" (mote than 30 widely dispersed 
localities). Brief ecological data arc Included where 
available, generally in the sequence: temperature 
°C), pH, dissolved oxygen (DO, mg 1 >), 
conductivity (Km, >iS cm ') and turbidity 
{nephelometric turbidity units <NTU)}, 

To minimise Ihe citalion of previous references 
to the rotifer families reviewed here, early Australian 
references which are included in Shiel & Koste (1979) 
are not repeated, nor are the majority of European 
references, which are included,, for example, in Ihe 
reviews or Hudson & Gossc (1886, 1889), Weber 
(1898), Voigt (1957), Bartos (1959), Rudescu (t9M), 
Kutikova (1970) and Koste (1978). A more detailed 
ueaunenr of the Rotifera outside Australia is given 
by these aulhors. 

Family Euchlanidae Bartos 
The following genera were integrated by Kemarte 
(1929-1933) in Ihe subfamily Brachionidae. A 
special family was erected by Bartos (1959), later 
accepted by Kutikova (1970) and KosLe (I97K). With 
the exception of Manfredium the genera are 
characterised by a lorica with plates which are 
connected with sulci, a segmented foot, more or less 
elongated toes, a corona of the same type 
i£nchtants-typt, Fig.3d) and mallcaix trophi 


W. KOSIt A R. j. stun 

K*y lu grnen* of the Family Kuehlanittae 

I I ■ r t ;i thlti without lateral sulct, long iocs 

« XtufiJrt'tJiuw Gallauhcr 

i orka rigid, With (fctfssl and sometimes virtual plate? 

2(1] Lurica composed of three platen, dorsal divided, 
separated by sulci ... . . Diptuis Gosse 
Dorsal pink- not divided 3 

JO), Dorsal plate arched concave; narrower than aiehed 

ventral plate , Dipfauchlams 

Doisal plate arched, convex or triangular 4 

4(3). Dorsal and ventinl plates nearly S same size, 
connected by a laieral double longitudinal membrane 
within a stiff' projection. . . . Tnpteuchtanis Myers 
With and without lateral longitudinal >uJc< 
i .Em'htunb Lhrcriberg 

Genus Manfrcdium Gallagher 
f-tnhwiyhtu Maufredi. 1927, p. 58 (non Wnlsiiuhartk 
Bcuuchampielta Kcmanc, 1929, p 107, 

Eudactylota Manfrcdi, 1927 is a junior homonym 
oi Eudactylota Walsingham. 1911 (Lcpidoptcra). 
Beauvharnptella Remane, 1929 is a nomen nudem 
under articles 12 and 16 of the International Code 
of Zoological Nomenclature (name published 
without description, definition). Manfredium was 
piopused by Gallagher (J957) to replace Eudactylota 
Manfrcdi. There is a single species, Manftvdiutn 

MunJrwJtum euducf viol urn (Gosse) 

Sctiridiurtt eudactvlutum Gosse In Hudson 2c Gosse^ 
1886, p. 74, Fig. 21:4. 

Eudactylota eudactylota (Uos-hc) MaulnoJt. 1927, p. 58. 
HcaUchumpwth euduclvfota (Gosse) Remane, 19*9, p, 

Man/radium eudaavlaww (Gossc) alter Callaghej. 

urn. p m. 

Diagnosis; Body pear-shaped with soft, 
transparent cuticle; head small; loot with two longer 
segments; toes long, rod-shaped; striated muscles 
visible in caudal part of body and foot; nvastax in- 
toned, tropin malleate (Kg. 1:1); oesophagus with 
small paired glands; large red cerebral eye; resting 
Ctf with fine hairs (Vtg. l:e r . 

Length; 420-760 >im, tout 30-110 am, toes 
275-320 /*m, (ropbi 25 am. longest uncus tooth 12 
/an, manubrium JS^nt. 

Distribution: Cosmopolitan, pancontinental In 
Auvualia, rare. Wannstcnothcrm. II.5-17,U°C\ pH 
4.8-6.5, conductivity to I60U pS cnr'. The rotifer 
may swim between water plants, also jumps by 
1'licking the roes. 

Literature: Kostc 1978, 1981: Riddei lVKLKotJc& 
Robertson 1983; Koste & Shiel 19S7a 

Genus Diploid Gpase 
ptpkfctkwk in Hudson & Gossc, 1886, Vol 2, p. 86. 

Diploic duv testae Gossc 
FIG, 2;I 

Dipiois davtesiue Gosse In Hudson and CjOffo IS86. 
Vol. 2, p. S7. Fig. 24:3. 

P, phlegraeu Iroso, 1910, p. 301. 

D. sculpturara naday, 1897, p. 153 (from hew Guinea). 
Diagnosis; Lorica with three plates; arched dorsal 
plate divided longitudinally by sulcus; lateral sulci 
small; toot l haw segmented; toes thin; lorica surface 
smooth, flexible; head short, broad; eye with leru; 
oesophagus short; stomach compact, ^tsiric glands 

Length: to 500 am, width 225 fan, toes 100 wo. 

D&trihulion; May be cosmopolitan in the bent hie 
of freshwater Sphagnum pools. Not yet recorded 
from the Neotropies. Not recorded in this study, but 
there is an early record from Qld (Thorpe 1887). 

Genus Dipkuchhnus Bcauchnmp 
Pipteuvhtum Beauchamp, 1910, p. 122. 
Dorsal plate narrower than ventral plate; tool 
three-segemenied; toes mote or less long, rod- 
shaped with tapering points occasionally swollen 
terminally; rami points with minute comb (Fig. 
2:2b); unci with 7-10 teeth; gastric glands long in 
younger individuals, broader and lobed in adult; 
red cerebral eye. Two variants of a single species (D. 
propatulai are known from Australia; individual 
can be discriminated by toe structure. D. propatuh 
has toes without a swelling before the points, 
whereas D. pmpatula nmcrodactyla has toes with 
a swelling. 

Diplcucldanis pmpatula propatuta (Gosse) 
FIG. 2:2a-F 
Diplois pivpatuUi Gosh- In Hudson & Gov,e, Ikflft, p 
87, Fig. 24;2. 

Dipleuchlanis pmpatula (Gosse) alter Heauchamp. 
1910. p. 122. 

Diagnosis; Caudal part of dorsal plate, sometimes 
stretched to rounded point and reaches nearly lo 
end of ventral plate; toes tapering to short point. 
Length: 338-508 ^m, lorica length to 500 jttHU 
width to 200/un; toes 70--I10 nm. 
Distribution: Cosmopolitan in littoral, sometimes 
occurs in plankton of ponds or billabongs, Rare, 
Vic, NT. Warm stenotherm, pH 4,5-6.4, also in ffce 
poor waters on the surlace of mud. 
t.itmttme: Koste IV7S, 1981. 

Dipleuchlanis proputulu macrodaavtu (Hauer) 

FIG, 2;2g 
Pipfeucfdatns tnavrvducty/a Hauer, 1965, p. 351. 
Dipleuchlanis prvpattdu f. macroihKtvtv (Hauer) altci 
Koste, 1978, p. J45. 

Diagnosis: Caudal part of dorsal lorica with shoit 
paired cuiicular projections; toes long. 

l-engtb: 12K-244 ,j,rn, width fvM-211 «m. toe* 
1 12-168 m . 



hig 1: Manjredmm eudactyiotum (Gosse): (a) lateral; (b) ventral; (c) dorsal; (d) lateral; (e) resting egg; (f) trophi; 
(g) opened rami; (h) uncus: (i) fulcrum. Fig. la-d, t'-i after Wulfert (1940); e after Kosrc (!978). s 'Scales lines, top 
100 pm (a-d); lower right 20 ptfa (e); lower left 10 /*m <i-i). 


W. KOSTii & R. J. SH1KL 

Fig. 2: I, Oipfois daviesiae Gos&e* dorsal. 2, Diptcuchlams propaiula pmpatuta (Gosse): (a) dorsal; (b) trophi; (c) 
ventral, swimming; (d) toe; (e) unci; (0 cross-section; (g) D. propatula macrodactyfa (Hauer), dorsal. 3, D elegans 
(Wierzejski): (a) dorsal; (b) lorica cross-section (ct O, propaiula). 4, Tripleuehldrtis plicata pficata (Levander): 
(a) lateral; (b) trophi; (c) dorsal, swimming; (d) cross-section; (e) dorsal; (f) ventral; (g) trophi, apical; (h) lateral. 
5, T. plicufu ruzeUni Rodewald: (a) lateral, (b) dorsal, swimming; (c) ventral. Fig. 2:1, after Weber (1898); 2a, b, 
f, 3a-d, alter Myers (1930); 2c t d t 3, after Fadccw (1924); 2g, 3c-g, after Hauer (1965); 5a-c, after Rodcwald-Rudcscu 
(I960). Scale lines Fig. 2:1-4, 100/iin (adult lorica each number group); 5,50 pan; 2b, 4b, g, 10 /an. 



Distribution Patttropical. NX, Vic Waler to 32°C, 
pi J 5.4-6 5 

littwtm: M*WS 1930; Haiter 1965; Koste 1974, 
m\\ Ber/aris 1982. 

Doubtful or insufficiently described species: 

Dipieuchtani.Ke{e%anspS r ierr.ejskl 1»93) = t&tchfotys 
prcpotulu efeftons {Fig. 2:3| according to Deauchamn, 
1910. p. 122. 
Dipleuchlatm COnradi livens,. 1947, p. 17<J, R* 5, 
Dipteuclitortte (KiludtKaHaviCT. r936.p. 139, Fie. 2:12. 

Genus Tripleuchfanis Myers- 

TcipieucMarits Myers, 1930 p. 379. 

U>rica ovoid in shape, truncate in front, 
smoothly-rounded caudally; dorsal and ventral plate 
nearly tyt same size, connected by pair of lateral 
longitudinal sulci. Longirudinal flajige of stiffened 
cuticle extending for entire length between each, 
giving bellows- like appearance to cross-section. R>ot 
thrvc-jolnied. guarded by shield-like process 
extending downwards from median longitudinal 
flange; toes short ending in abrupt points; rnasrax 
of modified malleate type (Fig. 2;4h); six club- 
shaped teeth on each uncus; rami triangular with 
minute denticulate combs at inside of tip; double 
cerebral eye, A single species, 71 plicate. w»lh a 
doubtful variani (T. plkaiQ f. ruzelmi (Fig. 2:5)) 
from Romania distinguished un I he basis of an 
elongated dorsal plate, caudally with a rounded 
notch (Rodewald 1940:88). 

Triptetwhtanis ptitwa (Lcvandcr) 
FIG. 2:4 

Euchlan'u; pitvaw Lcvaitder. I8<M p. 4X. 

Tripleut :hlun f& pfkuta t Levanderi after Myers, 1930, p. 

Diagnosis: Lorica stout; foot glands and reservoirs 
elongated; retroccrebral organ small, 

length; 250-270 jhb, dorsal plate 9(M30 ,«m. 
ventral plate 100-115 /mi, toes 25-37 txm. 
Distribution; Cosmopolitan; rare in fresh-, more in 
brackish and marine waters, alfiO In warm springs 
QId T Vic 
t-iterature: Hauer 1925, Kostc 1978. 

Genus Euchhviis Ehicnbcre 

Eucbfonis Etirenberj;, 1832, p IM. 

Lorica Iranspaient with oval or ovate outline; 
dot sal plate arched, convex, sometimes with median 
keel or lateral wings, larger than ventral plate, 
variable in height and *liape; venrral plate caudally 
somewhat indented,, nearly flat. Two plates 
connected by thin, flexible cuticle forming 
longitudinal sulci. Foot with two or three joints; toes 
long or short, sword-shaped or parallel-sided, with 
sharp tips. Pajted tojig seUe situated on dorso-distal 
purtlo-n of foot-segments (Fig. 3a;ss). Corona of 

family type (Fig. 3d); long sensory bristles and cilia 
in tufts atid solitary on apical field, Trophi (FS&. 
3e f i) modified malleate; fulcrum <Fig. 3f:fjl broad, 
pointed rami (ra) with and wirhoui minute comb; 
uncus with main and variable number of accessory 
teeth, Brain with large cerebral eye (Fl$. 3b ey) 
behind large mastax. Large rclrocerebral organ 
present (Fig. 3b;re), with subcerebral glands. Dorsal 
antenna large; lateral antenna tFig. 3b:la) with 
sensillae tufts on small tubules. Stomach .sometimes 
with sacculi but not constant between species. 
Excretory organ with large contractile bladder (Fig. 
3a,b:bl). VitcJlarium with eight large nuclei, Male 
thinly loricate < Fig. 3I:m) with reduced alimentary 
tract; proionephridia present. Hx>i with two to three 
joints; two long sensory bristles on terminal joint. 
Toes short. Dark resting eggs may be attached to 
plants (Fig. 3hj). 

Euehfom species are littoral rotifers living among 
aquatic plants, but in the pclagial they occur in 
cu trophic lakes and ponds at Cyanophyta maxima* 
especially of Gtueotriehia, the main food of 
Euchfanis dilatata (Ruttncr-Koti.sko 1974). Other 
'rinds consists pf diatoms, dexmids, other algae and 

The taxonomy of the different species is difficult; 
even within ihe same population lorica shape and 
cross-section i.s variable. The vfwpe of the anterior 
margin of boih plates i< of little value in the 
determination. Most useful is trophi structure; 
which is species-specific. Eucktanis should not be 
pressed by a eoverslip. See for example & fftencta 
crtHs-seeciuns and the different forms of £. dtlatcra. 

Key *« Species of the Genus Euchfanis 

I. Cross station of lorica more or less arched to 

semi-circular 2 

Crow section of lorica triangular, dorsal plate vefa 
high keel \q 

2(1). Posterior ed&e ot dorsal toxica wirh distinct notch 

or enibaynienl (cf. Fig. 7:1k. l-tg. 8;2a). 6 

Notch ahsem, or only shattow wmargination tefi Wg*. 
4:3b. fe.U " ...3 

3(2). Vemral plate present 4 

Venter membranous or rudimentary - 5 

413) Ventral plate ca. i& dorsal ptaie width; wing-Ill^ 
lateral expansion* of dorsal lorica niargin (Fig, 4:3b) 

E. </ lata Vononfccw 

Ventral plate ca. Yi dorsal plate widrh, constricted 
al puuerior end; in> expanded margins iFie. 6:3) 
E. tyro Hudson 

5(*). lateral eoustrictionM'n medial dorsal lorica; flanged 
lateral margin* l.lig, 8;3> ..£. pynformti Gwse 
Dorsal touca moi constricted, literal margin not 
flanged (FifiS 4:3; 4:4) £ dtflexa Gossc 

6|2) <_ utJeuIar shield-like process just below caudal part 

of dorvil plate (Fig. 7.U) £ meneia Mver* 

Shield-like process lacking. . . '. .7 

7(6 1 Ventral plate rudimentary; longitudinal sulci absent. 
ioet. very long (>!00 jam), stender <Fie, 4;2) 

* • - £• Q$tjpif8a Myen 

Ventml plan, well developed . h 



hie. V Euchlnms dilutaiti ritlataia Chrenbery: (a) ventral; (b) lateral; (c) cross-section; (d) apical field; (e) tropin; 
(f) trophi, lateral; {%) k. triquetra, amictic Efgi (h) mictic egg; (i) E. dtluhtta, mictic egg.: (jl subitaneous egg; (k) 
sperm; (i) male; (m) E. deflexa, male. iKey to a,b: bl = bladder; dpi = dorsal plate of Iorica; cy =■ eye; o,g = 
gawc glands; la = lateral antenna; m - mnmh; me ? membrane; ov = ovary; re = jcirocercbral sac; si = 
stomach; s^ = sensorybristles; vi = vitellarium; vpl - ventral plate of loi'ica. Key to f: ma = manubrium; fr 
= fulcrum; ra = ramus; u = uncus). Fig. 3a, c, j, after Kosle (1978): b, after Bcauehamp Qm$)\ d. after Rematie 
(19 U); e, after Hauer (1930); f. after Voigr (1957); g, h, after Wesenberg-Luud (1929); i, al'tci Kozar (1914); k, after 
Ruttner-Kolisko (1972); 1, after Wulfert (1956); m, after Lcissling (1924). Scales lines, top, a-d, g-m, 50^m; e, f, 10 ( tm. 

8(7). Dorsal Iorica with winglike lateral extensions 
deflected vent rally at tips (Fig. 8:2)/:. phryne Myers 
No lateral extensions of dorsal Iorica 9 

9(8). Foot and iocs slender: rocs [ A length of dorsal plate, 
blade-like, fusiform (Fiji. 5;lb)£. tiilatuia Ehienberg 
Foot and toes stouL; loes >4 length of dorsal plate 

toes- dilated di.stally for vk length and constrict to 

acute points (Fig. 8:1) E. orophci Gosse 

lG(l).Venlral plale coinpletely developed, lateral sulci 

present (Fig. 6:1) « E. mcisa Carlin 

Ventral plate reduced to thin hyaline membrane; wide 

dorsal plate flanges (Fig. 8:Ie-e E. triqueiru 




Euchlunis alata Voronkov 
FIG. 4:1 

tzucManis alarti Voronkov, 1912, p, 210. Figs 2*3. 
Diagnosis: Shape of body ovoid: dorsal plate 
without terminal notch or embaymenu bvit may 
have lateral vving-Jike extensions (see Fig, 4;lb, dL 
Wingless individuals cosily mistaken for £. lyra or 
£. deflexQ. Intermediate forms common, Ventral 
plate elongate, relatively narrow, but without 
constriction before end as in £. tyro. loot two* 
segmented and robust; toes Sloul and fusiform. Tips 
of rami incurved with pair of finely denticulate 
combs; lour stout teeth in each uncus with several 
accessory teeth. 

Length of dorsal plate 260-36Q jun; ventral plate 
width 102-156 ^m; toes 70-9g^m; toes width 12-20 

Distribution- Nearctic and PaJeaxctic, in acid boreal 
waters,, littoral and psammal. Not yet recorded from 

Literature; Pcjler 1962; Kostc J978; Chenjsalath & 
koste W3. 

Euchianis calpidia Myers 
FIG. 4:2 

Fucklams calpidia Myers, 1930, p 37] , PI. 20. Figs t-S. 
Diagnosis. Dorsal plate round, or highly domed 
("obscure triradiate", i.e. helmet-like) in cross 
section. Middle portions of lateral edges straight 
or pinched, extending downward below venter; 
ventral plate rudimentary <cf. E, (kffaxa and E. 
pyftformis), rudiments connected wuh dorsal plate 
by flexible membrane; lateral sulci absent; foot two- 
jointed, slender; toes very long, parallel-sided wilh 
abrupt point. Tropin characterised by five slender 
paired teeth in unci with iwo or three acccssoiy teeth 
beside smallest. Tips of rami without minute 
denticulate comb. Tropin similar to those of E 
dfflexa and E pyriformis. E, calpidio distinguished 
by presence of distinct posterior noich at end of 
dorsal plate, and by relatively long toes. 

Length of dorsal plate 280-400 //in: lorica width 
220-397 pnv, toes 120—155 /an; toe width to IS uttj. 
Distribution: Nearctic- Paleqrctic I Europe 10 bar 
fcasi). Single record from a GouJburu R. hitlabong 
near Seymour, Vie. 22°C^ pH 7,3, DO 8,0 mg \-\ 
Literature: Bauer 1936; Wang 1%l;Kutilcova 1970; 
KOsle 1978. 

Euchianis deftexa Gossc 
FIG, 4: 3 
EuMents deftesa, 1K51, p 200. 
Paptdte (kflew (G&uft in Myers 1950, p. 3t>9. K. 21, 
Figs 1-5. 

tDapidia = EuMants afier C*r(Wi ( VW, p. 16) 
lynxgaosis; Body resembles arc of circle In cross- 
section; dorsal plate ovoid, without distinct 

posterior notch, sometimes wilh shallow 
emargination. Lateral edges of dorsal plate 
connected by flexible membrane somewhat 
Ihickened in position occupied by ventral plate in 
other Euchfanis. Posterior portion of this area more 
hardened as rudimentary ventral plate. Nu 
longhudinal sulci present. Foot stout and *wo- 
jointed, long setae on dorsal end of first joint; 
tropin wilh five long teeth on each uncus, clubbed 
at tips, with 2-3 accessory teelh; rami drawn out 
to long lips, minute inside comb* lacking <Fig. 4:30. 
Unglh of dorsal plate 19G-350 ftm t width 
140-240 ^m, toes 55-100 /mi, toes to J5 ^oi wide, 
subltanenus egg 180x89 ^di. 
Distribution' Cosmopolitan in ihe littoral, 
occasionally in the pelagiaj. Rare; NSW., fas., Vic, 
W.A. There may be local variants, e.g. the 
Tasmanian foiiu closely resembles E. deftexa targa 
{big, 4;4), described from Lake Balkasch, Siberia 
(Kulikova 1959). !4.0-21(J*C , pH 6,0-7,7, do 
8.0-8.6 mg I"', <70fiScni | 
f iterature: Evans I95I; Koste 1 978; ShieJ & Koste 

Euchtants ditatata Ehretibcrg 
FIGS 3. 5:1 

huenhnrs ditatalu Ehrenherg, 1832, p 131, Fig 4:3. 
Diagnosis; General body shape ovoid, truncate 
anteriorly, rounded posteriorly; dorsal plate variable 
in height and shape of cross-section. Varying fioin 
low arc ol circle to high triangle; ventral plate 
almost as large as dorsal; longitudinal sulci not deep 
and narrow (Fig. 5;U), Posterior dorsal plaie 
divided by deep elongate notch of variable form. 
ct f. hwksianu (Fig, 5:4), f, untsetuta (Fig. 5:3) or 
f. largo (Fig. 5:5). Foot slender and two-jointed; 
paired sciac on distal margin of fiisi segment; toes 
mo.%tJ> parallel-sided, tapering terminally io sharp 
lip. Unci of trophi with four opposing teeth; ventral 
large teeth each have rudimentary tooth; tips of 
larni with inner minute denticulate comb; 
retroccrcbral sac large (Fig, 3b;re, cf. also Fig. 5:41}; 
brain wilh red eye on dorsal side. 

Iiilraspecil'ic forms ate described exclusively 
according to differences in the shape of the lorica 
and cross sections, however research by Parisc (1965) 
suggests that different morphological types are 
correlated with ecological differences. This also may 
apply to some distinguished species; E. parro 
Rousselet, for example, resembles E, d'thtata and 
is distinguished from it by smaller <jze and lung 
slender toes. All known measurements fall within 
the range of variation of E. dtiaiata; it is seen as 
an ecotype of £. diialuta. 

Lorica length 140-320 >tm, dorsal plate width 
100-255 f im, central plate 95-172 /im, toes 50-100 
am, posterior notch 12-61 ^u deep; male 115-148 





^ 4b " 4c Jf - 4d v 

Fig 5: I, Euchlanis ditatata dilatata Fhrenberg: (a) dorsal; (b) toe; (c) cross-section. . £ ( dilatata mavrura (Ehrenbcrg): 
ventral- 3, E: dilatata f. unisetatit (Leydig): (a) dorsal; (b) cross-section; (c) toe. 4, E. dilatata f. lucksiana (Hauer)r 
(a) cross-section; (b) dorsal; (c) ventral; (d) trophi, apical; (e) swimming; (f-g, k) cross-sections; (h, )) different 
lorica forms; (i) e. dilatata, lateral, swimming. 5, E. dilatata larga Kutikova: (a) dorsal; (b) toe. Fig: I;a-c, 2, 3a-c, 
5a, b, after Kutikova (1970); 4a-d, j, k, after Hauer (1930); 4c-i, after Myers (1930). Scale lines 100 fctJL 

Fig. 4: L Euchlanis alata Voronkov: (a) without wings, ventral; (b) with wings, ventral; (c) toe; (d) ventral; (c) habit, 
lateral; (0 cross-section. 2, E. cafpidia (Myers): (a) dorsal, swimming; (b) trophi, apical; (e-f) cross-sections; (a) 
lorica, ventral. 3, E. deflexa Gosse; (a) dorsal; (b) lorica, ventral; (c, g, j), cross-sections; (d) trophi, apical; (e) 
toe, lateral; (f) other trophi; (h) dorsal; (i) resting egg; (k) lorica, broad form, ventral; (I) loe. 4, E. deflexa fargu 
(Kutikova): (a) lorica, ventral; (b) toe. Fig. 4:1a. d-f, afier Yamamoto (1953); lb, c. 3j, 1, 4a, b, after Kutikova (1970); 
2, 3a-e after Myers (1930); 3f-h, after Donner (1964). Scales lines 100/*m (adults lorica in each number group). 



j<m, toes 34-240 #nt [E, pttrvo Rousselet, IS92; 
dorsal plate length J40 ptt, width 100 ^m, ventral 
plate length 125 /an, width 75 /an, toes 70 //in, depih 
ttl' posterior notch 50 ^m|. 
Distribution: C Cosmopolitan in fresh and brackish 
water: pH 4-10, CI 140 g I" 1 (Koste 1978). The 
most common and widely tolerant cuchlanid in 
inland waters; pancontincnlal, including Tasmania. 
8.0-29.9°C, pH 5.44-7.9, DO 3.6-13.8 mg I -\ 
23-161K) f i$ aer\ 2-235 NTU. 
titetvlure: Myers 1930; Hauer 1915; Kurikova t970; 
Ridder 1972; Koste 1978: Shiel & Koste 1979; Koste 
£ Shiel 1980. 

Euchtants ixcisti Carlin 
FIG. 6:1 

EwMoms inctsu Carlin, \93>9, p> IT. 

Euchlams trigueira Hudson &, Gotse, 1839. PI. 23, RjR 


Dtagnosis: Body ovoid in. shape, truncate in front 
with variable anterior notch; dorsal plate i/iradiale 
in cross-section with median keel extending from 
neck to an^le of poster lot v-shaped notch. Ventral 
plate completely developed, Longitudinal deep sulci 
connect both plates; toot two-jointed, setae on first 
foot joint; toes slender, fusiform; trophi (Fig. 6:le, 
b) with five teeth on each uncus. First and last (Fig. 
felly have accessory shorter tooth; minute combs 
on rami tips. Male (Fig. 6;li) with normal anatomy 
lor genus (cf. Fig. 3:1). 

Length of dorsal plate 210-270 ^m, ventral plate 
ttfKMO ptu. lorica width to 240 j*m, toes 70-85 *m; 
male 145 /an, toe* 26 /an. 
Distribution: Cosmopolitan in the littoral* 
occasionally m the pelagial of shallow ponds and 
biUabongvRare, NT., fas., Vic: 13.5-29.9°C, pH 
5.V7.V, DO 5.45-8.2 mg \~\ 23.0-80.2 #S em 1 , 
2.0 NTU. 

Comment: A variant, £1 tnquetnt mucronota i~ 
C incisit mucronaia) (Fig. 6:2a, h) described by 
Ahlstrom (1934) from the Ncotropics may be an 
ecotype; it resembles E ittetia and may co-occui. 
The dorsal plate has an elongated keel. Length of 
dorsal plate io 320 /uii, lorica width lo 252 (M\ 
height io 120 ,itn; toes to 120 /<m. Not yet recorded 
from Australia. 

Literature; Myers 1930; Wfelftrl '956; Kusie 1974. 
I97K, 1981; Smcl & Koste l<TOj Berzins 1982. 

Eucftiamt lyru Hudson 
FIG. 6;3a-j 
Much/tints tvra Hudson In Hudson & Guiic. 1886, i\ 
8% Pig ??:!. 
£* rnyersi Kutikova, 153.9, p. 223. 

Diagnosis: Body elongate and ovoid; cans-section 
r^einbles arc of circle; lateral sulci present; dorsal 
plate without posterior notch; wniral plate totally 

developed, somewhat constricted caudally; foot 
slender, two-jointcd v paired long. setae on fust foot 
joint; Uophi with five paired unequal teeth, wilh 
two or three accessories; rami tips with minute 
inside comb, Length of toes variable. 

Length of doisal plate to 335 /irn, ventral plale 
302 /*m, lorica width 110-180 /on, toes 76-90 /im. 
toe width 14-16 urn. 

Distribution: Cosmopolitan in inundation areas 
also in periphyton, Single record from L. Mulwala. 
Vic. 12.0°C, pH 7.5, DO 10.7 mg I" 1 , 46 }iS enf, 
17.5 NTU. Variants are known in the northern 
hemisphere, e.g. E. lyra myersi (Kutikova 1959; 223) 
(fig. 6:4) a Paleaicttc subspecies wilh long slender 
toes (86-107 /j.m long, 7-8 /mi wide) and E, lyra 
larga (Fig. 6:3k, I) (Kutikova 1959) from N. Siberia. 
Valiants arc not recorded from Australia. 
Literature: Myers J930; Pejlcr 1962; Koste W%\ 
Shiel 3c Koste 1979. 

Euchlunfc menefa Myers 
FIGS 7:1,2 
Euchlanvt mctwia Myers, 1930, p, 378, PI. 19. Figs 5-J* 
Hucfthuiis otopha Lucks, 1912, p. 105, Fij;. 31. 
Euchlanis proximu Myers, 19)0, r> 377. PL 19. 1-igA 1-4. 

Diagnosis.- Shape nearly ovoid; well-developed 
ventral plate almost as large as dorsal plate; cross- 
section semicircular arched but also roughly 
trir3diatc; dorso-vemral width of lateral sulci 
(relatively) uniformly wider iltan in any other 
congener; cuticular shiekl-like process beneath 
posterior notch above first foot joint; posterior 
notch of dorsal plate very deep and often large (Fig, 
7*.lk); foot two- jointed; toes very long and blender,. 
sometimes with minuie claw at tips. Male 
colourless, with short toes; dorsal and ventral plate 
still conspicuous; brain lone with red eye; two 
sensory hairs on liny tubules in apical fidd> as in 
female. Trophi (Fig. ~:le) with four main iccih, 
accessories not visible; minute denticutaie combs 
on inside of rami tips (Tigs 7*le» 2d). 

Length of dorsal plate 105-173 un\, width 80-M0 
/an, depth of posterior notch 35-45 /mi, width Of 
ventral plate 50-90 /an, toes 60-75 /*rn> width of 
toes 3.5 ;«m, Male total length 170 am, toes 25 /mr. 
Disinbauon: Cosmopolitan, mostly in acidic 
waters: rare in submerged Sfthagnvm; N, America, 
W. liftman*. N.S.W., NT. Vic.. W.A. 10.0-29.9°(\ 
pH 5.3-7.4. DO 5.45-10.1 me I \ 2(M23 /iS cm V 
22 NTU. alk, 2.6-2,7 m^ L ■. 
Literature: Myers 2930; Hauer 1935 (firoximp); 
WuIIctI 1951; Pfcjler 1962; Berzins 1963. 1982; Koste 
1978, 1981; Koste & Shiel 1980. 

Euchlanis oropha Gosse 

fk;. &i 

huchlunls ntOfttitr Ciwsse, HR7, p, 5, F-ig 2:16, 

nun Euihkmh vrophu LucKb, 1912. p. 10^, ng, 31. 



Diagnosis; Resembles E dilamta but usually 
smaller, with stouter foot and more robust, 
differently-shaped fuslfoi*m toes; latetal antennae 
situated in cavities beside somewhat keeled end of 
dorsal plate; irophi construction as£ dilafata (cf. 
figs 3e, 5:4d). Comb-like processes on inside of 
each ramus tip; four opposed club-shaped 
functional tccth\ 

Length of dorsai plate 164-266 am, widrh 
127—172 /tm, ventral plate length 123-225 T i<m, toes 
49-7H /<m» width 10-14 ftm, posterior notch 25-29 
Distribution: Cosmopolitan NT., QUI, Vic. 
8.0-18.5°C, pH 4,8-8.4, DO K 4-10,0 rH| H, 
67-400 **S cm ', 4.5-160 NTU 
Litervture: Mvers 1930. Kutikova 1959; Kcxste 1978, 
Shiel & Kostc 1979' Bcrzins 1982, 

Euchlanis pofomenew Bcrzins 
FIG. 7:3 

Euvhlatris parametria Uerzina, 1973. p, 126, Tigs- 4,5, 
Diagnosis; Resembles E, mmeta (see above). Iocs 
with pseudoclaws: anterior margin of dorsal plate 
wkli two short keels. 

Length of dorsal lorica 95-104 ^m; ventral plate 
70-72 m\ width, dorsal 7l)-75 ,tm, ventral 50-55 
>im; anterior margin width 33 /an, caudal opening 
26-30 pm long x 18-25 /oil wide; toes 45-50 /i.m. 
Distribution; Recorded from New Zealand* not yet 
known from Australia. 

Euchlanis phrvne Myers 
FIG. 8:2 
Euchlams ptuvoe Mycn*, 1930, p. 372, PI, 14, fig: fa 
l J l IS, Ftgs |-4 

Diagnosis; Dorsal lorica oval, with slight 
constriction of median lateral margin; deep inverted 
U-shaped caudal notch almosi as long as toes; 
stillened ventral plate connected to dorsal plate by 
longitudinal sulci; toes short, fusiform, ea. '4 length 
of dorsal plate; trophi with four club-shaped teeth 
on each ramus; rami with fan-shaped denticulate 
combs on inner distal margins. 
A superficial resemblance to E, pyriformis and E. 
tatpidia was noted by Myers (1930), and Kosie 
U978) considered E. phryne a possible variant of 
E. valpldiu or e. dilutatu. Comparison of lorica 
morphology of the species above, particularly cross- 
sections, and differences in trophi structure, 
indicates that E. phryne is a distinct species. 

Length of dorsal plate 225 pan, ventral plate 210 
um, dorsal width 180 /on, ventral plate width 150 
j*m, toes 65 ;*m. 

Distribution: North America (Maine); single 
Locality, a blllabong of rhe Ooulburn R. near 
Seymour. ViCi, in Spring 1 97ft and again in summer 
1978. 17,0-22.a*C. pH 7.1-7.4, DO 8.7-9,8 mg I \ 

Comment: The above anomalous distribution is 

noteworthy; E. phryne is one of seveTai rotifer an<i 
mierocruitaceau species recorded from Goulburn 
RiveT hillabongs with similar disjunct distributions 
(Shid J976 and unpublished data; Koste 1979). The 
importation of heavy dam-building machinery by 
the Utah Construction Company in the mid-1950's 
(during the construction of Eildon Dam) ts 
implicated a5 a possible source of transporr iff 
resting eggs/cphippia. Populations have become 
established in sheltered billabongs downstream o'' 
rhe dam site, 

EucbiantK pyriformis Gosse 
IIG 8:3 
Luihlams pyriformis Gosst, 1851, p 201 
Dopi'li a piriformis Myers, Uwft p. 370, Kig. 15:5-3. 
Diagnosis. Shape nearly circular, may have slight 
ernarejnaiions instead of caudal notch; lateral 
margins turned downwards, occasionally pinched 
in the middle; ventral plate rudimentary, only 
developed posteriorly; no lateral sulci; foot two- 
jointed; two long pairs of setae on dorsal side of 
the first foot segment; toes relatively short; trophi 
resembles that of E. deflexa (cf. Fig 4:3f). 

Lenglh of dorsal plate 285-320 /<m, width 
275-315 m< toes 80-85 /on, toe width 10-13 ^m. 
Distribution; Cosmopolitan »n the litloral, 
occasionally pelagial. Vic, Tas. 
Literature: Kosie I97K; Her/ins 1982; Shtd<& Kosic 

Euchlanis triquefv EJuenbcre 
FIG. 9:1 

Euchfamstri/jai'tralLriTcnberfr, 1833. 0.461, fig. ?7;g. 

E. pellucnii! Hiuring, 1V21. p. 6, Tig* 2. 

ti tatftubopi'tca Manfred i, 1927. p. 24, lig. 7K 

Oaptfiia cahnata Carlw-Vilssun, 1934, p- *. Via 2 

P, lata Cartin, I9J9, p. 16. 

Euchlanis truwerra pterig&ideo Crete, 1955, p 6(1, tl$& 
15. 19. 

non-/T. trh/wrtv frfter Ruitiier-KohsUi, IV72, p, |82, Fig. 

Diagnosis. Body hi dorsal view circular, sometimes 

somewhat elongated (Fig. 9:1); triangular with a 

high keel in cross-section; median keel extends from 

Jtcck to caudal region; dorsal plate laterally with 

wide flanges (Fig. 9:te-e>; no ventral plate, nn 

posterior notch, no lateral longiiudinal sulci; some 

specimens with semicircular ventral line over base 

of toes; foot obscurely two-jointed; toes long, 

slender, neatly straight, ending in abrupt points, 

most lorica* very transparent with coloured innei 

organs (retrocerebral sac dark, mastax yellow, 

stomach olive green, yellow or brownish) Male i Fig. 

9;3!i,t) with cuticular plates and dorsal keel. Trophi 

cf Fig, 9:lj-U| Ups ol rami with minute comb* (Fi^. 





Lorica length 30O-710 um, width to 540 /im, toes ornamented with distinct pattern and cavities; toe* \hort 

90-15 /on, lorica height Lu 300 ,*m. trophr rumi 40 lophocharis Ehrenberg 

...m, fiilcrum 32 ,<m, mamibTia 50 jan. Genus Mvtriino Bory de St Vincent 

Dtsinhutioti: Cosmopolitan in the Utioral, M yii Una Bory dc St Vincent 1826. p. 87 \*$etpwa 

sometimes pelagic in shallow waters, N,S t W.» QM, Khtenherg, IK30, p. 46 = Oipia.\ Gossc, 1851, p. 201 = 

NT., Vic. Oiplaadium Lauterborn, 19H, p, 4K3). 

literature.- FVrjIcr 1962; Kostc 1978, 3981; Shirt & Two recognized form-scries ("Formenkreise"). 

Kostc 1979; Benin* 1982. One series urongly loricate, usually with variable 

Stiff anterior and posterior spines; anterior margin 

T-, r ii j of lories generallv granulated; toes stiff and sword- 

The following species are not considered here: . yZ*_T^* X , , • 

r- \* knit i .if ki a shaped second eroup thin v oncate without 

E. Myers, 19.16, known only from N. America ' . . . - . . ~ ,« . 

and ?Volga; anterior spines (except At ucartthophom, (Fig. 10.1, 

& bioculatu Use!, 1901 {nomcn tmdem name published 2); anterior margin not granulate, toes mostly long 

without description, and apparently later synonymiacd and slightly curved* flexible* 
wuh E. plkata Levander by Isset ( 1 *>f )fi >. Incorrectly cued 

m r™VTl\ * nd ISffil!!Si -,,••■ Key to Species of the Genus Mrtitin* 

E LVitWti Mvctk I'MOfHg. 9;4) impossibly a juveride t • . 

form of E. triquetra, Only known from the Nearctic '* lrtr,c f . tTollt ' anterior margin gianulated, ItrjW 

Length of dorsal plate 170 ^m, lorica width 80 «m\ toes s*orcl-shaped, . , ,,,,, .2 

Mi/*m. (Myers, I9"K>; Kostc, 3078); Lorica thin, anterior margin not granulated, iocs 

E coUimorpha Berzins, 1957 (Fig. 9:2) resembles £ i t% Ia "S and . curved . . . . , J 

intita, only known from Gambia. W, Africa. W. Anterior lonea mat gin with two shot! dorsal and 

E, contorts Wulfcn, 1939. known only from central l *° ^° r f vcnlra ' srnne3, Xl f,WCf0flata MAIlec 

German* and the Volg^ csmary; Anterior tnnca margin with only two ventral spines 

£. dupidulu Parisc, 1966, seekoMe I97H fmisquorcd as « • , ' • ' : * -. - . M . ****$* (Ehrenbtrg> 

Herise, 1963). Doubtful species; ™) ventral margin oi lorica with variable projections 

£. hvpMdavtvto Hum ' 19GJ, see Kusle 197* .Doubtful •;••-• -. W jWftf^W* Hauct 

species. Ventral anterior margin without projections. . .'4 

E ui>ulata Kuttkova A Vas^lcva, 1WZ. endemic in I ate *3l J oe * w ! 1 ' 1 cJaw ? 5 

Baikal^ Toe* without claws, . ,.6 

t*. mamorokaensis Her2ins, N73 (Fig. 9:3), recorded SW tle 3d pari loricate, 3 fool segments 

from Madagascar; - .• A/, crasstpes I Lucks! 

£. mikmpous, Koch Althaus, 1963; recorded only from Htfad P aTI "lorieaie; 2 toot segments 

central Germany; £t V"*"," . . . ***' ""WW® fLucks) 

fi patwomco Baruch, 1ST?, known only from Hungarv, n(4 '- SS reiat|vc,y shorl; *«« 'J»"Pa W lor '" ^t" 

single record; loldi A7. bisulatta LucKsj 

£ pvrpusUUi Ridder. 1977. known only frmn -the Toes long, sti;uelit, anleriur margin wilhout folds 

Caribbean; pttedsptaa Si.\tcanthophora).M. trigona (Gosse) 

£ ( Riidpw-j Id . 1 940>. know n only from . „ .. . , , _ 

Romania Myhhna ocanthophora Haucr Myt.lmulae BartOS Mmlwaacwhophoru Haucr, J^i p. 550, Fi &S 73a-c 

fflSfSS 7 0SS h SeC H° nS 0t ,0 ['1 2*8 *&** dorsal lorica M^ngl arched, 

ZBS t! J? r rh , on ; blc ^ emM ! o!ate "J gmnuiated, ventral lorica flattened, with depression 

dorsolateral pla es firm y fused: lone dorsum with E XilZJ ■ i i • . , 

,. , i ,. r, . beneath head openine; tnant?u ar lonca proiecrions 

or without sulcus, latter common with double Keel; nZl L^^JSyS^L ^ . ?iL Va^S 

.tl, uTifc.,, iJlt. t :. . , , ' rlanK medioventral head aperture (T is* lOrlbfc 

three or less tout sections; ocs pointed, straight or u ...i • .. ' .. '* 

.i.rrh.u, .„.«,- 4 ,».,».,i i ii I? am shield-hke projection over two-segmented toot; loes 

vliehily curved vcntralwards; malleate iropht. AIJ r on y nointcd 

spedes littoral and benthic, occasionally (but rarely) r' «.k, IS >«^ „/..!. >«a ji - ^ 

in ( h« plankton. Two genera (see Bartos #5 h J?, 8 o C W 5 ^oT^ 

Kuhkova 1970, Ko,« e 1978). J?* IO g «« w ' d,h t0 64 '; m: T ^T? 4 ^ ' 

' An unusually large specimen from Lago Cirande, 

ttw .- > ^# ».* --is -j Amazonja had the following measuremcfii& 

Key |o ecnera of Mylil.nidae (sequence as above): U2, 198, 125. 137. »4Q-l44 W f 

1.1 oricathm or ncid with dorsal sulcus and double keel; f r ncfp linrt ,.Kt \ 

lonoa nnoniamented; toes Ions (Koste unpubl ). 

Mytilina Bory de St Vincent Distribution: Probably panlropical in polysaprobic 

Lorica without dorsal sulcus, one strong keel; lorica waters. Not ye! recorded from Africa. Occasionally 

Hu. fi: I, }Zut:hton*\ tm:i\<j Cartin: (a, h) lorica cross-jectious; (c) venlral; (di lateral, swimming; ie| tmphi, apical; 
(I) dorsal; <ut toe; On) uncus; (i) male. 2. E t/icisa mucronata (Ahlstrom): fa) dorsal; (b) lateial, 3» E tyra Hudson 
(a) dorsal; lb) venlral; (c) trophi, apical; (d, c) cross-sections; if) dorsal* swimming; fg. h) lonca. ventral; Ii) mc; 
(j) toe; (k) E. tyra (orga (Kutikova), ventral; (1) toe. 4, E. tyra myerst (Kutikova): (a) ventrah Ihi another form, 
veil nal; (c) tlQphi, apical; (d> crosi-tecrion; (e) toe. Fig, 6: la-t. 3a, b, 4a-d, Myers (1930); if, u, 3c-l, 4c, after 
Kutikova (1970); 2a, b, alter KoMe (1974). Scales lines 100 ^m (adult loiicd in each numbet group). 


\V. KOSTE& K. J. SH1H. 

Fig. 7: 1, Euvhianis meneta Myers: (a) lateral; (b) dorsal; (c) ventral; (d) lorica cross-section; (e) trophi apical; (f) 
lorica ventral; (g) cross -section; (h) lateral, swimming; (i) lorica dorsal; (j) toe; (k) dorsal., swimming; (!) dorsal; 
(m) cross-section; (n) maJe; (o) male, lateral 2, E. proxitna Myers; (a) lateral, swimming; (b) cross-section; (c) ventral; 
(d) trophi. apical; (e) dorsal: (f) ventral: (g) dorsal. 3, E. purumeneta Ber/ins: (a) dorsal; (b) ventral. Fig. 7:la-e, 
2e-g, after Hauer (1935); If, g, i, j, after Kuiikova (1970); Ik, o, after Wultcri (I960); 3a, b, after Beams (1973); 
2a-d after Myers (1930). Scale lines, top, 100 /an (Fig 7:1, 2), bottom, 50 urn (3a, b). 



in plankton samples. Qld (Townsville, Mt Isa). 
Literature: Rome l*>78; Shiel & Koste 1985, 

Mytifirw bisulcata (Lucks) 
FIG. I0;3 

thphix bisulcata lucks, 1912 p. 95, Figs. itSHL 

At rrigana alter Hairing, 1913 p. 75. 
Diagnosis: I.orica with (bids on anterior margin; 
three weak dorsal keels in cross-section; sulcus 
indistinct; toes shorter than in M. unguipes with 
long tips (cL M. unguipes (Fig. 1 2:2)) 

length 130- 1 HO m, toes 60-70 jtm, tips 8-14 jim. 
Distribution: In peat bogs, areas of decomposition. 
NX, Vic. 

Litentturvr Hau<r 1936; Koste 1978; Kosie&Shiel 
1980: Bcrziiu 1982; Tail et ai 1984. 

Mytiltna crassipes (Lucks) 
HG. 10:4 

Dip/ax crassipes Lucks, 1912, p. 96, Kig. 30. 

Mytititut cfiwipe* after Carlin-NHsson, 1934, p. 11. 
Diagnosis: Lorica laterally compressed; head wilh 
two lateral lounded plates; trophi small* uncus with 
<ix teeth; mouth opening with palpar organs; 
stomach and gastric glands large; two black ->pols 
at "forehead" region (?eyes). 

l-ength 220-280 *<m (total); lorica 138 fiov, toes 
57-75 fiXRZ claw 9 /an; tropin 21-30 p.n\. 
Distribution: Cosmopolitan in polysaprobic waters, 
occasionally in shallow-water plankton. NX 
(Magela Ck). 

literature Koste 1979, 1981; Kostc & Shicl 1980; 
Tait et al 1984. 

Mytilina mucronata (Mullet) 
PJQ. 11:1 

Brachionus imtcromtus Multer, 1773, p. 134 
Mytilina tnuirumilu niter Hofstcn, 1909. p. 54. 
Sec Koste (1978) for extensive synonymy. 

Diagnosis: Shape Erf lorica variable,, of tittle 
taxonomjc value (eg. M. mueronata var. spirtigera 
(Lhrenberg, 1832)); corona as in Luchlanix\ lorica 
oranulaied anteriorly dorsaUy curved spines visible 
in lateral view beside loot opening; keels cud in 
short hook; foot- three-segmented; largt cerebral eye 
present (Fig. ll;la eye lateral .antennae indistinct; 
dorsal antenna (Fig ll:la da) visible; trophi of 
malleaie type, unci with five teeth; mastax (Fig 11:1a 
rns) with salivary glands; stomach cellular with 
small gastric glands (fig. 11:1a gg). Male 
approximately half female sire, also loricate with 
ceiebral eye (cf. (Fig. 3:1, m)). 

Length:' 170-250 *tm, height 96-100 /on. toes 
53-60 |iiu, Subitaueous egg SO /mi, male lo 135 offl. 
Distribution: Cosmopolitan in eurrophic waters, 
3tso on the surface of decomposition areas between 
water plants and in brackish waters. Occasionally 
in the plankton of shallow ponds and billabongs. 

Vic. !3.5-22,0°C. pH 7.1-7.4, DO 4.1-10-2 nag \-\ 

240 pS cnr\ 5 NTU. 

Literature: Evans 5951; Koste 1978. 

Myubna tri$vnu (Gosse) 
FIG. 12:1 
Dtptax trigttrra Co.«t 1851, p. 201. 

Mytfiinu irixona after Hariing, 1913, p. 75. 

Diagnosis: Surface of lorica slightly dotted; 
anterior margin with rounded edge vent rally (cf. M 
acanthophora Fig, I0;l, 2); cross seeiion almost 
tiiangular; narrow doubk keel; loes with long tips. 

Total length: 245-265 ;int, lorica length ] 5(3-160 
/<m, toes 68-83 pJSL 

Distribution: Cosmopolitan m decompose- 
lion /inundation areas, Qld. 
Literature: Kosie 1978: ShielA Knsie 1979; Bcrzins 

Mytilina unguipes (Lucks.) 
FIG. 12:2 

D'tplax unguipw Lucks 1912 p. 96. 

Mytilimi bkukvta ./.' Uftguipcs (Lucks) after Cailhi- 
Nils*cm, 1934, p. 1L 

Mytilina ungmpes (Lucks) after Wiszniewski, 1953. p, 

Diagnosis: Anterior margin of lorica without folds, 

laterally compressed; loot with two joints; loes lung, 

slighlly curved with distinct, inflexible claw. 

Doubtful species, possibly identical with M. 


Length 193, height 94 *m, toes 58 ^m. claw 

II /itn, 

Distribution: European peat bog, nor yel recorded 

ftom Australia. 

Literature: kutikova 1970; Koste 1978 

Mytilina ven t talis (Chrcnben?) 
Was 11:3, 4, 5 

Satpima vcmralts fcluenberu. 1832, p. 133, PL 4, Tig. 7. 

Mynhna ventrutis (Ehrenberg) after Harrtnt, 1913. p. 

For cxtcnMve tynwiymv sec Harring tfoc. at.) and 
Kuukova (1970:519-520). 

Diagnosis: Lorica margin granulated, shape and si/c 

of lorica and spines very variable; only vcniryl 

anterior spines. Short-spined form (var. brevispina, 

176-215 /ml) shown in Pig, ll:3a-c; long-spmed 

form (var. macracantha, 250-350 ^m) shown in Fig, 

11:4. Fig. 1 1:5 shows rare form described by Wulfert 

(1965) (A/, centralis f. longidactyla) with short 

anterior but long median and shorter caudal vcnti&l 

hooks, Lorica (90-115, nn) without spines, median 

caudal spine 62 ^m, ventral caudal spines 48 jim. 

loes 72 /*.rn. These forms may be ecotypes. 

Distribution: Cosmopolitan. Probably 

paticontinental. not yet recorded from S. (f. 

typ from N.S.W., N.L, QUI Tas., Vic; brevispbw 

from NX, Qld, Vic; macracantha from NT, Qld, 





W.A ]3.5-2S.5°C pH 5.44-7.9, DO i.S-lOjB mg 

\'\ 23-1500 ti$ cnv ', 6.8-67 NTU, alkalinity 2-7 

Not yet recorded from Australia: 

M. hiearinuta (f'erty, 1850) 

Af, matniLfm (tenninns, 1894) 

Af. mw//«r (Hetty, 1849) 

M vitiens (1-evander, 1894) 
(wc Kosre W7H for details). 

Genus Lophocharis Ehrcnbrrg 

I ophocharis Khrenberg 1838, p. 458. 

Strong lorica, surface soft, granulated, but may 
have honeycomb structure; cross section nearly 
rhomboid but specimens with flexible integument 
nearly oval or compressed; dorsum always arched; 
more or less distinct keel with and without 
transverse folds; ventral plate with longitudinal and 
transverse ribs; apical margin with and without 
serration; ventral aperture rounded, dorsal more 
triangular; foot-opening ventral; Foot with three 
joints, toes with sharp points; uophi malleate (Fig, 
IV Id); cerebral eyes only visible in living animals. 
Littoral and benthic in habik seldom found in the 
plankton. Seven species are known, four have been 
recorded from Australia 

Km I* SpwUs of lhf Genus L^prWikxr/* 

I. Lonca ^120 pm long with distinct dorsal Keel 

producing triangular cross-section Z 

Lorica <100tffii lung, keel indistinct, cross-sccnon 
more tlliplicaJ, . . jL curvaio Heroins 

2(11 Anterior margin ol lonca strongly serrated, dorsal 
keel with transverse foluSA. salpinu U'hrefibere) 
Anterior margin smooth or lightly serrated, dorsal 
keel without folds 1 

?(2l. Lorica smooth, dorsal notch Of bend ape' lure V- 
shaped, ventral a broader L -stupe; fine striae on 

both nicies of dorsal Y.eti /.. naias Wullen 

Lorica nulpiurtd as J., salptna, head aperture 

notches Y-sliaped. no striae beside keel. . .£. 

oxysternoti (Gosse) 

Lophocharis curvata Berzins 

fjg. tsa 

Lophocharis eurvata Berrtns. ls*K2, p. \Z, Fig. I5a-c. 
Diagnosis: Small species, lorica elongated U-shapc 
without typical dorsal keel, cross section more 
dorso-ventrally flattened than m congeners; head 
aperture margin with dorsal and ventral sinuses, 
ventral slightly broader; foot aperture broadest 
posteriorly; toes thin, cutved outwards. 

Lorica length 95 jwm, width 55 /*m, toes 18 /*m, 
fool apetture width 32 /mi. 

Distribution: Endemic, single record from King 
Parrot Clc, Kinglake, Vic. I8.X.53. 
Comment: Although the original description and 
figures of this rotilet do not meet the requirements 
of the Code, we feel that the distinctive dorsum, 
foot apermie and small size warrant retention of 
specific status until material can be examined. 

Lophocharis naias Wulferi 
FIG. 13:4 
Utphoehw is tunas Wuifcn, 1942, p. 1WS, Fig. la fc. 
Diagnosis: Dorsal lorica with shallow keei and 
lateral longitudinal striarions, surface morphology 
Otherwise smooth, unornamented; apical margin of 
lonca finely serrated; uncus with 7 teeth, inner 
margin of rami with fine denticles. 
Lorica length to 170 pin, width to 105 ^m, toes to 
21 pim. 

Distribution: Europe. Single record, from Coongie 
Lakes, S. Aust., coll. Jane Roberts, Botany Dept, 
Univ. of Adelaide, 06.xh.86. 
Literature; Kosle 1978, 

Lophocharis oxysternon (Gosse) 

FIG". 13:1 
Metoptdia oxysternoa Gosse. 1K51, p 201 
Lophocharis oxvsternun (Gosse) after Harting, 1916, 
p, 564, Fig, 97:6-13. 

Diagnosis: Surface of integument soft or weakly 
pustulated; ventral plate with deep depression over 
foot-opening. Many intermediate forms to L. 
salpinx e.g. with slightly serrated apical margin, 
Trophi (Pig. J3:ld-c) with si\ uncus teeth. 
Lorica length 120-200 ;im, toes 24 /±m. 

Distribution: Cosmopolitan, benthic in fresh ard 
brackish waters* occasionally in plankton. Vic. 
Literature: Koste 1978, Berlins 1982. 

Lophocharis salpinu (Ehrenberg) 
FIG. 13:2 

LvpadeHa salpina Hhnenherg, 18.14. p. 209 

Metopidia satptna after Hudson & Gosse, 1S89, p, 46, 
Fig. 34:4, 

Lophocharis salpum (Ehrenberci after Harrine, 1916. 
K 563. 

For extensive synonymy see Kostc (1978:15.1) 
Diagnosis: Anterior margin always serrated; keel 
with transverse folds mostly over caudal part of 
lorica; variable, intermediate forms to A, 
oxysternon; trophi also with six uncus teeth; foot 
with three joints; toes sharply pointed, vent tall v 

Length 175-224 fun, lonca 120-135 >un; width 

Fie 8: I, Euchiunis oropha Gosse: (a) dorsal; (b) dorsal; (c) cross-seel ion; idt alternate caudal lorica*; {ej tropin, 
apical, 2, £. phnne Myers; (a) dorsal; fh) ventral; (c) trophi, apical; (d) cross-section. 3* £ pyr(formt\ Odsrc; (A) 
ventral (b) lonca umss-seei ion: (e) rrophi, 4, t', riSysM Myers: (a) lateral; (t>) cross-section; (c)irnpbi: (dl ventral 
R& 8:la. alter Koste (W8); t-d, alter Donner (\9Uy, le, 2a-d, for*, 4a-d. after Myers (1930). Scale lines 100 
atn (fig. 8:1-3); 50 /<m (4a, b, d). 



Fig. 9: 1, Euchlanb triquetra Hhrenberg, (a) lorica, ventral; (b) lateral; (c-c) different cross-sections; (f) dorsal, swimming; 
(g) toe; (h) male, contracted; (i) male, lateral, swimming; (j) trophi (fulcrum and rami); (k) ramus; (1) manubrium; 
(m) fulcrum; (n) uncus. 2, E. callimorpha Berzins, dorsal. 3, E. mamorokaensis Berzins: (a) dorsal; (b) cross-section; 
(c) ventral, 4 ? E. perpusiUa Ridder, ventral- Fig, 9:1a» after Kutikova (1970); lb-e, after Grese (1955); lf-n % after 
Wulfert (1956); 2, 3a-c, after Berlins (1973); 4, after Riddcr (1977). Scale lines 100 ,tm (adult lorica in each number 



3a 3b 

Fig. 10: 1, Mytilina acanthophora Hauer from Magcla Ck, NX: (a) ventral; (b) lateral; (c) cross-section; (d) lateral 
(cf. 3a, b). 4, M. crassipes (Lucks): (a) lateral; (h) dorsal; fc) lateral; (d) dorsal; (e) lorica, contracted; (f) trophi; 
(g) trophi, lateral; (h) uncus; (i) mouth with palpar organs (m = mouth). Fie. 10:3» after Hauer (1936); 4, after 
Wulfert (1939). Scale lines 100 /*m (adult lorica in each number group). 






Fig, 11: 1, Mylilina mucronata(0. F. Miiller): (a) swimming, taLeral [/?/ = bladder, 6/- = brain, rf<? = dorsal antenna, 
eye = cerebral eye,./' = foot, gg « gastric gland, / = intestine, wht = mastax, &t = stomach, t= toe, vi = viLellarium]; 
(b) male, swimming, lateral; (c) lorica, lateral; (d) lorica, dorsal; 2, Mytilina sp. cross-section. 3, M. ventralis 
(Ehrenbcrg): (a) lorica, lateral; (b) ventral; (c) male, dorsal. 4, M. ventralis macracantha (Gosse) lorica, dorsal. 
5, M. ventralis longidactyla CWuKeil), lateral, from E. Africa. Fig. 11:1a, 2, 3a, b> 4, after Kosle (1978); lb, c, after 
Pejler (1962); Id, 3c, after Weber (1898); 5, after Wulfert (1%5). Scale lines 100 pm* Top Fig. ll:lb-d; left, 3, 4; 
bottom, 5. 



Fig, 12: 1, Mytilma trigona (Gosse): (a) lateral; (b) ventral; 
(c) lorica, dorsal; (d) cross-section; (e) swimming, lateral; 
(0 ventral. 2. M. unguipes (Lucks): fa) lateral; (b) cross- 
section. Fig. 12:1, after Donner (1954); 2, after Hauer 
(1936). Scale lines 100 p,m. 

80-95 /*m, toes 25-40 /tm. 

Distribution: Cosmopolitan, benthic in fresh and 

brackish waters, also in polysaprobic pools and 

ponds. Pancontinental. 10.5-30.4°C, pH 5.6-8.4, 

DO 3.7-11.0 mg \~\ 28-3330 /*S cm -\ 5-92 NTU, 

alkalinity 4.0 mg I '. 

Literature: Evans 1951; Shiel & Koste 1979; Koste 

1981; Berzins 1982. 

Not yet recorded from Australia; 

Lophocharis hutchinsoni Edmondson, 1935 

L, naias f. ambidenta Dc Ridder, 1960 

L. parva Rudeseu, 1960 

L rubens Wulfert, 1939 
(see Koste 1978 for details). 

Family Trichotriidae Bartos 1959 
Three genera with loricate head and body; surface 
with facets, mostly granulated and with spicules or 
spines, sec. on dorsum; foot freely movable or with 
stiff joints; trophi maJleate. Generally occur 
between aquatic macrophytes and in periphyton. In 
plankton only as migrants. 

l'ig. 13; 1, Lophocharis oxysternon (Gosse): (a) dorsal; (b) ventral; (c) lateral; (d) tropin, apical; (e) trophi, lateral 
2. L sulpirw (Ehrcnbcrg), (a) swimming, lateral; (b) lorica, lateral; (c) laieral; (d) another lorica, lateral; (e) cross- 
section. 3, L. curvata Berzins, (a) lorica, dorsal; (b) cross seclion; (c) lorica, ventral. 4, L. naias Wulfert, (a) tonca 
dorsal, (b) lorica vcntraL Fig, 13:1 a-e, 2b, after Harring (1916); 2a. after Wulfert (1956); Id, 2c, after Haucr(I92S> : 
2d, after Klement (I960); 3 after Berzins (1982); Scale lines, centre, 100 ,im (la-c, 2-4); top right, 10 ^m (d e) 


\v kostp & r. j. SHIPl 

Fig. t4: Wolgaspinijpro (Western): (a) dorsul, *witninnig; 
<b) lateral; (c) venirai; (d) cross-section a-c after 
Western (1894); d, after Vranovsky (1957). Scale line, 
50 fira. 

Key to genera of Trieboiriidae 

I Lorka with anal segment - 

Lorica withoor anal segment, . . . tyotga SlGDftKO^ 

2(1). Lorica with distinct elnngated spines on 

Mocmchoews Ftrty 

No long spine* on loflcft 

Trichvttia Bory do St Vincent 

Genus Wolga Skorilcov 
Wolua Skorikov, 1903, p. 37. 
Monospecific, p$0y0 spintjt'ra. 

WtolfiQ spimfera (Western) 

FIG. 14 

Ohtyfu winifent Western. 1894, p. 427. Fig. 21:1-4. 
Cathypna spintfera after Murray, 1913, p, 545-554. 
/.ecane spinifera after Harring, 1913, p. 62. 
Wotga spimjeru (Western) after Skoiikov, 1903, p. 37, 

Diagnosis: Dorsal lorica divided by ribs into panels 
or facets; foot noi loricate! freely movable; ventral 
plate with oval foot-opcuing under Transverse 
distinct line; toes short, slender, pointed. Cross- 
section of body somewhat compressed (Hg. 14d). 
Male unknown. 

Tbtal length 200-255 /im, lorica 95-145 /tm, width 
70-90 /*m, spicules 5-6 jim. 
Distribution: Cosmopolitan In fresh and brackish 
water, raicly tfl ihc pelagic. Two localities, probably 
more widespread, but rare. N.S.W., Tas 10.5-1 IXPO, 
pH 6.7-8.4, DO 10.0 mg 1 \ 
Literature: Koste 1978, Koste & Shiel 1980. 

Genus Trkhotria Bory dc St Vincent 
Tmhowui Bory dc St vincenl, 1927, p 752. 
Head, body and three foot segments, heavily 

loricate; cross-section hexagonal (Fig. IMfJ (sec 
comment with T. buchtwri for exception); inosl 
lorica borders with spicules, greater on "shoulders* 
if head contracted (Fig. 15:3, 4a); lorica with facets. 
granulated; head lorica composed of several plates' 
second foot segment with teladvely long spines (Fig. 
1 5:1a fsp)\ toes long; corona of Euchlanis type; 
tropin (Fig. 15:Id-e) malcate; red cerebral eye (Fig. 
15:1a eye). Male loricate (Fig. 15;lb-c), Food algae 
and detrilu.s. diatoms preferred. Trichotria occur* 
rarely in the plankton (as migrants). Seven species 
recognized, with many variants; five known from 

Key to Species of (he Genus Trkhotria 

I. Cross-section hexagonal, three foot segments.. 2 
Clttss-setuon triangular, two foot-segments 

77 bachneri Kosie et oi, 

2(1). Only distal end of terminal foot segment and to*.-. 
projecting beyond posterior lorica margin; lorica 
elongated U -shape. . .'/: pseudocurtu Koste et at, 
AiJ ihree, or at leasi two, fool segments *nd iocs 
projecting beyond lorica margin ....3 

3(2). Last toot joint with dorsal mimire spine between 

the toe bases t pocitlum (MulleO 

No minute ipine between the toe bases 2 

4(3)- Lutcial pail of doisal lorica conspicuously large; 
marginal spicules of lorica direcied forward 

T. truncaia Whitelegge 

Lateral lorica not very expanded* spicules if present 
not directed forward. . . T (etracits (Fhrenberg) 

Trichotria buchneri Koste, Shiel &, Tan 
FIG 15:1 
Tfichotno buchneri Koste, Shiel & fan, 1988, pp. 
120-121, Figs 3-5. 

Diagnosis: Lorica of triangular cross section with 
median keel ending in elongated caudal spine; 
dorsal lorica with longitudinal rows ol denticles to 
keel (appears granulated by light microscopy); 
anterior dorsal margin deep U-shaped aperture 
between two lateral cusps; ventral maTgin with a 
shallower U-shaped aperture projects beyond dorsal 
margin to level of lateral cusps (or slightly beyond 
lips); rectangular plates cover head,, wben 
contracted a pyramidal projection extends to or p3ust 
anterior lorica margins; foot two-segmented. 
without spines; toes long, rigid, with acute points 
Length 1 60-182 fifty width to 125 <un; height to 
84 /(.m; proximal loot segment 24 ^im, distal 15 /*m; 
toes 50 jtm. 

Distribution: Known only from marginal reed beds 
of acid dune lakes north of Strahan, on west coast 
of Tasmania ().. Garcia, L. Ashwood and 
surrounding dune pools). 17.0-20.0*0, pH 3.1-5.15 t 
8().6-IXH.4Vs cm \ 0.9 NTU 
Comment: The morphological differences between 
this species and congeners were noted by Koste et 
ul, (1988), particularly the domed and keeled cross- 
section, caudal spine and two-segmented foot. The 



Fig. 15: 1. Trichotna huchneri Koste et ai a. dorsal, b. lateral. Cl ventral; 2. Trtchotria pseudocurta Koste et ai a 
dorsal; b. ventral. After Koste et ai (1988). Scale line 50 pun. 

armoured head is peculiarly trichotriid amongst the type locality in Jan, 1989, and additional sites for 

Rotifcra, but the distinctive features of T. buchneri the rotifer recorded. A detailed SEM study of this 

suggest that generic status may be appropriate. species will appear at a later date (ShicI & Koste, 

Accordingly, additional collections were taken at the in prep.). 



°, e eye 



Triehotria pocillum (Multer) 
FIG. 16:2 

Trichoda pocillum Muller, 1776. p. 281. 
Dinocharis poctllum after Chrenhcro,, 1830. p. 47. 
Ttkhoiria pociUum Dory de St Vinecm, 1827. p. 7.52. 
For extensive .synonymy \ee tlarrinc <I9)3| and Koste 

Diagnosis: Length of font segments and spine on 
second Fcoi segment variable; lorica spindle-shaped. 
surface divided into granulated facets by ridges, 
corona simple, can be retracted in neck and lateral 
head plates: male loricate Rare in plankton. 

Length 200-300 gm (total), lorica length 110-140 
}i\\\ foot spine 55-66 ^m, median minute spine on 
last loot segment 20-28 pm, toes 83-145 Mm. 
Distribution; Cosmopolitan in the littoral of fresh 
and brackish waters. N.S.W., Qld. 
Literature: Shiel & Koste 1979, 

Trichotria pseudocurta Koste, Shiel & Tan 
FIG. I5;2 
INehotttQ paeudocurlu Koste ct at 1988, pp. 121-3, Fie. 

Diagnosis: Lorica small, U-shaped in outline with 
both surfaces granulated; triangular cusps at 
external angles; two raised ribs on ventTal surface 
terminate anterior to raised rim of foot opening 
similar ribs form twin keel on dorsal surface, 
commence either side of median notch on dorsal 
anterior margin, fuse to a single keel on posterior 
lorica; foot 3-segmentcd, without .spines; toes with 
short claws 

length to 123 >4m, width to 74 ^m; loesio 30*mi. 
Distribution: Known only from hunnc waters of 
Lake Pedder (White Spur Inlet), Tasmania. 2I°C, 
pH 5.2, 32.6 fib enr\ 0.5 NTU. Resembles T. 
curta (Skorikow) from the Volga R.. however the 
latter is smaller (110 pm long) with more angular 
morphology, and lacks the frontal cusps. 

Triehotria tetraelis (Hhrenherg) 
FIG. 16:1. 3 
Dmocham U'iractis Ehrenberg, 1830, p. 47, 
trichoma tenarris (Thicnbergt after Carlin 199$ p. 40 
For extensive synonymy sec Koste (J*>7$:157). 
Diagnosis: All morphological characteristics 
variable; lorica spinulaled or not; measurements of 
lorica, foot segments, dorsar and ventral lorica 
ornamentation differ between individual*, with 

morphological series between 7! truneuta and T. 
tetryetis similis (latter has very long toes; 120-190 
/mi <Fi.e 17)); caudal pari of lorica with one or 
paired hooks (T tet metis eaudata; ipinulcs on 
second foot segment may be veiy* short (T, tet metis 

Length typical form; 218-380 /<m, toes to 160 fflfy 
T tetraetis simiits (Stenroos, 1898): 330-400 ^m> 
second foot segment 42-4K /un, loes 120-190 ^m; 
T tetraetis eaudata: 230-260 /mi, caudal hook of 
lorica 22-28 jim, toes 50-72 pm. 
Distribution: Cosmopolitan in macrophyies. 
littoral. Commonly in shallow waters of billabongs 
in N.S.W,, Qld, Vic, (typical form) and NX 
(stmilis), J3.5-:4.5°( , pH 6.3-8.0, DO 5.8-9.2 mg 
\\ 59-400 ^S cm \ 2-160 NTU. 
Literature: Koste 1998, 1981, 19,S4; Shiel & Koste 
1979; Green 1981; Ifer/ins 1982 

Trichotria truneaia Whitekgge 
FIG. I6:4a-b 
Dinacharis trumarum Wlmelegge. 1889. p. 315. 
Dirhu'haris intermedia Bcrgcndal, J892, p. 107, Tig. 
Trichotria cornuta Myers. 1938, p 11 
Trtehatrfti teltwOs var- truneaia (Whitelegge) in Koste, 
1978, p. 1S7. 

Diagnosis: Margin of lorica with spicules, also on 
foot segments, all directed forward; most with 
shorter toes as in T tetraeiis, but man) intermediate 
forms; variable facetcation on lorica. 

Length 95-146 run, grealest width l2S;*m, loes 
60-10(1 ;tm, 

Distribution: Acidophil, (H Sphagnum* mainly 
between plants, seldom in open water. N.&.W., NX, 
Vic. 25.0-28.5°C, pH 5.44-7.6, DO 6,0-6.2 mg I' * 
23-145 >iS enr 1. 

Literature- Koste 197S, 1981, 1984. 
Comment; Although morphologically variable, T. 
truncarum is readily distinguished from T. tetractis 
in material we have examined from both Australian 
and Canadian collections (cl. Oiengalath St Kobte 
1983|, and wc consider it a distinct species. 

Not yet recorded from Australia 

/ mrarw raudara (I uek$ t lWi;cf Ko»te I9&4, p. 116; 

/ teuctctis pauperu (hhrenberg, 1630); 

T. curta iSkorikov, 1914), 

T Mtkowcta Myers, 1934, 
(See Koste. 1978, pp. 157-8 for detail*!. 

fig. 16:1, Triehotria tetractis (Ehrcnuetg); (a) female, lateral, swimming; \a = anus, ht - hladder, hr => hram, dt 
= dorsal lorica, da = dorsal antenna, eve - cerebral eye; ,4' = fool eland;,/) = foot segment;/*/; = loot spine*. 
gg = gasnic gland, ta ~ lateral antenna, oe = oesophagus sr = salivary gland, 47 = stomach, vi * vitellarium, 
vt = ventral lorica]; (b) male [do ■ doisal antenna, eye - cerebral eye,/ - foot, p - penis, pg = prostate gland, 
ss * testis, vft = ve^iigial gut], (c) male, lateral [ef = CtDHKd fields, coiona» eye — oei'diial eye, p = pems, ss 
= reRiis, v$ = vestigial gut]; (a) trophi \ra ~ rauiu 1 ;, w = uncus nra - uutmibiium]; fc) uncus, latcrai; (T) lorica, 
cross-section, 2, T. pociUum (Mullcr), female, dorsal, jwinimiri^; J, T, tetractis, contracted ttoJWf. 4, T. truncaia 
(Whitelegge): (a) dorsfll, from Canada; (b) from Romnnia 177 truttcata t, hnzispma after Rodewale) I935| ftg. 
I5:l», ), 2. }, 4rt Jfier Koste |i9***K 4b, alter Rodesvuld (l!?*?5). Seak hnes. leu 50 /mil (Hg J5*Ja-cj; lowei rijjltt. 
100 Mm {2-41 







/ mm 1 
3b f ] 

lpftl ? 



Genus Macrochaetus Perty 
Lorica flat, hexagonal cross-section, head part 
retractile; lorica granulated with minute spinules; 
more or less developed anal section; two loricate 
foot segments; toes generally spindle-shaped; paired 
symmetrically inserted spines (Fig. 18:1a); lorica 
margin with spicules, shoulder hooks larger (Fig. 
18:1a, 7a-b); integument opaque; corona simple, 
with central buccal field and longer lateral sensory 
hairs (Fig. 18:3b); dorsal and lateral antennae on 
visible papillae; one cerebral eye; mastax with 
malleate trophi; eggs and males unknown. AH 
species live between or on water plants and are rare 
in open water. 

Key lo Species of the Genus Macrochaetus 

[refer to FIG. 18:1a] 

1. Only one pair of rudimentary anteriosubmedian 
spines present (2a, b).M. danneeli Koste & Shiel 
More spines present 2 

2(1). Two anterolateral spines, two anteriomedian spines, 
two posteriomedian spines present, central dorsal 

spine pair absent (4a, b) M. cotlinsi (Gosse) 

Central dorsal spine pair present 3 

3(2). Two anterolateral- , two anteriomedian- , two 
central median- and two anteriosubmedian spines 

present M. altamirai (Arevalo) 

Two anterolateral-, two anteriomedian, two 
anteriosubmedian- and two posteriomedian spines 
present M. subquadratus (Perty) 

Macrochaetus altamirai (Arevalo) 
FIG. 18:3a, b 

Polyehaetus altamirai Arevalo, 1918, p. 1-47. 

Macrochaetus esthonicus Riikoja, 1925, p. 7, Fig. 2j. 

Macrochaetus attamirai (Arevalo) after Selga, 1952. 

M. austratiensis Berzins, 1982, p. 18, Figs. 17, 38. 
Diagnosis: Shape, size and spicules at lorica margin 
variable; distinguished from M. co/linsi only by 
paired central dorsal spines (see key to species). 

Lorica length 80-115 /mi, width 76-108 /mi, toes 
20-28 ixm. 

Distribution: Cosmopolitan warm stenotherm, rare. 
Probably more widely distributed in Australia than 
a single record from Victoria would indicate (as M. 
australiensis by Berzins 1982). 
Literature: Koste 1978; Shiel & Koste 1985. 

Macrochaetus cotlinsi (Gosse) 
FIG. 18:2 

Dinocharis cotlinsi Gosse, 1867, p. 269, Figs. 1-4. 
Macrochaetus cotlinsi (Gosse) after Harring, 1913, p. 67. 
For extensive synonymy see Koste, 1978, p. 161. 

Fig. 17: Trichotha telractis similis, lateral (toes 190 /un). 
After Koste (1984). 

Diagnosis: Shape, size, length and number of 
spinules on margin variable; number of dorsal 
spines constant; lorica brownish. 

Total length 94-250 /mi, lorica 53-112 /mi, width 
62-112 /mi, second foot segment 20-25 /mi, toes 
16-20 /mi, dorsal spines 48-66 /mi, posteromedian 
spines 24-48 /mi, anal segment spines 32-35 /mi. 
Distribution: Pantropical and subtropical, warm 
stenotherm, benthic. NT., Qld, Tas. 28.5-29.9°C, 
pH 5.44-5.47, DO 5.45-6.2mg 1\ 23-44 /tS 
cm -1 , alkalinity 2.6-2.7 mg l\ 
Literature: Koste 1978; Shiel & Koste 1979; Tait et 
al. 1984. 

Macrochaetus danneeli Koste & Shiel 

FIG. 18:5 

Macrochaetus danneeli Koste & Shiel, 1983, p. 116, Figs. 

Diagnosis: Lorica granulated/pustulated; dorsal 
lorica with terraced sides to blunt keel; margin of 
keel base with longer spinules; uppermost keel 
dorsum covered with large cuticular ledges and 
beading; anal segment spineless; shoulder spines 
(Fig. 18:1a, 7a), posterolateral- (5a, b), 

Fig. 18: 1 (a) points of insertion of spines and setules in Macrochaetus Perty spp., dorsal lorica and foot segments 
\la, b = anterolateral spines, 2a, b = anteriosubmedian spines, 3a, b = anteriomedian spines, 4a, b = central 
dorsal spine pair, 5a, b = posterolateral spines, 6a, b = posteriomedian spines, 7a, b = shoulder' hooks, 8a, b 
= anal segment spines, 9a,b = foot segment]; (b) Macrochaetus lorica, cross section. 2, M. collinsi (Gosse): (a) 
dorsal, swimming; (b) lorica, ventral. 3, M. attamirai (Arevalo): (a) dorsal; (b) ventral. 4, M. subquadratus Perty 
(a) dorsal; (b) dorsal; (c) ventral; (d) lateral. 5, M. danneeli Koste & Shiel: (a) dorsal; (b) ventral Fig 18ia b 
2a, b, 4a, 5a, b, after Koste & Shiel (1983); 3a, b, after Rodewald (1940); 4a-d, after Wulfert (1964). Scale lines 
50 fim each number group. 




posteromedian- (6a, b) and only one pair of 
rudimentary anterosubmedian spines present. 

Lorica length 125-130 /an, width 130-137 pm, 
foot segment 20 /*m, toes 16-18 pm. 
Distribution: Single locality known: Buffalo 
Billabong, Magela Creek, NX 30.9°C, pH 5.38, DO 
4.15 mg \\ 29 /*S cm" 1 . 

Macrochaetus subquadratus Perty 
FIG. 18:4a-c 

Macrochaetus subquadratus Perty, 1850, p. 22. 

Potychaetus subquadratus Perty, 1852, p, 45, Fig, 1:6. 
Diagnosis: Lorica generally hexagonal (Fig. 18:4a); 
ten dorsal spines of variable size and shape; 
shoulder spines most conspicuously large. 

Lorica length 80-135 /on, width 73-140 /wn, 
second foot segment 8-16 /an, toes 15-22 fin. 
Distribution: Cosmopolitan in periphyton of 
aquatic macrophytes. Rare migrant in plankton, 
more common in billabongs. Thermophile. 
Literature: Koste 1978; Shiel & Koste 1979; Green 

Not yet recorded from Australia 

M. hauerianus Wulfert 1964 (-M. coilinsi van Hauer 

1938), p. 534 (Indonesia); 

M. longipes Myers 1934, p. 13; see Koste 1978 p. 161; 

M. multispinosus Myers 1934, p. 12; recorded from the 

neotropics and S. Nearctic; 

M. phitopax Wulfert 1961, p. 87; single find in central 


M, sericus (Thorpe) 1893, p. 182; see Koste 1972, Koste 

& Shiel 1983 p. 117 Fig. 8. 


The support of the Australian Biological 
Resources Study in grants to RJS, the use of the 
facilities of the Botany Department, University of 
Adelaide in manuscript preparation and long-term 
loan of microscope facilities to WK by the 
Deutschen Forschungsgemeinschaft are gratefully 
acknowledged. The Department of Lands, Parks 
and Wildlife, Hobart, supported fieldwork in the 
World Heritage Area in Jan. 1989. Particular thanks 
to the staff of the Peter Till Laboratory, Albury- 
Wodonga Development Corporation (now Murray- 
Darling Freshwater Research Centre) for providing 
collections from billabongs in their study area, to 
Professor D. G. Frey, Biology Department, Indiana 
University for the donation of an extensive series 
of rotifer publications from his Limnology Library, 
and to an anonymous referee for critical comments 
on an earlier draft. Our thanks also to Charles 
Hussey (BM(NH)) for checking obscure references, 
Anne Grant, CS1RO Publications Service, for 
editorial expertise, and Lor Wai Tan for final 


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and iophochurts with descriptions of live rtew species. 

i*roc, US. Nat, Mus, 51, 527-567 

.11921) RotulOiw. Repi. Caned Arctic Expert 

097&1 Rotatoria Oie Radcritere Miileieuropas. 

Bcstiinniunubwerk bcgrundel von Max Voigt. 2 VoK. 
fBornlraeger, Stuttgart t 

U979) New Rortfcra from the River .\«uxr*v, 

IW-1918, Ottawa. 8. £1-21 
HuifcH, J. (1925) Rouitcincn aas den Salzgewasseni von 

Oldesloe (HoUtein). Mitt Geagr, Ges. Nat. Hist, Mm. 

Lubceli II 30, 152-195- 
0930) Zur Roi3lorienlauna DcutschJand* (1). 

7oo/. (Jfla 92, 219-222- 
(1935) Zur Rotaionenfauna Outschlands (IV), 

ZooL Anz. 110. 260-264. 
. .0936) Raderttcr aus dem NntuischtM^gebiei 

Weingarrencr Moor. Ret?*, nutarkdl Forsch. 

SudwestdeutschL KartsruHe 1, J29-I52. 
(1938) Die Roratorien von Sumatra, Java und Bali 

ttach dfif Efftcbniss^n ovi Deutsche^ Liitniotot;isch<:ir 

Sunday Expedition. Arch, ftvdrobtot, Suppt, '5* 


.(1965; Zur Roiaumen fauna des Aina/oiuis^ehieies. 

Int HeU gfifi Hydrubiot. SO, 341-89. 
HonrrtN, N. vo*j 0909) Rotatorien aus- dern Kiastermyr 

(Onrtfand) und einijjen andern .strnvedi'ichen 

RtnnengewaViern, Ark //W, <j, 1-125, 
Hudson, C. T. & G<J&6, P. H. (1886) The Roiifera or 

whecl-ammalciilcs, both British and foreign', 2 vols. 

(Longmans. London). 
<1S89) The Rotifera Or wheel-animalcule?., both 

British and foreign*. SuppL (Longmans, I on don) 
IROSO, l t (1910) Pritrto cOnlribufa aJla conoscen-ca At'i 

Rntiferi dt-l laso-stagno crarerico di Astroni, Matt, Zont, 

Hoi 21. 299-"304 
Ksn R (1901) Btiufi miIIr Jernwlc cutanea. i.Nota 

prcveniiva). Boll, Mus, Z06i. Uiiv. Genova IflH. 5 pp. 
^^___ (lyOot Sulla termobior,! negti animah aquarici. 

RtccTvrn? iaunKtichee^iiVc^o//c. Ait Soc, Ligtixstica 

Soil Nat Gfttova 17, 5-72. 
JcNNtMusII.S. (1894) A list of theRotaiotiaof the Great 

Lakes and some ofrhe mland lake; of Michigan. Bud. 

Mich. Fish. Comm. 3. I-3J 
Kicmc^t V 096(1) 2ut Routoneni'auiu ephemerer 

CieW'asser //;, Vtr VQjttt, VuturktU; Wu'ruenihtf^ 115, 

K*.( h Ai fHAUs. B. (1962) Weiiere «eue und 

hemerVcnswci te Kut^itonen aUs dem StechltiHec 

Umnologtca Heel I, 65-85. 
KosrE, W, (1972) Rotatorien aus Gewasvern Arnazonicns. 

AtWtZOtitiiM 5. 25H-5D3. 
(I974» Roia»t>nen aiii eincm L'ler»ecdes unLeren 

Rio Tapajos dem Lago Varotn ^Amii/orvien f. &cumsx. 

Abates, 53. 4V6«. 

iOLitheaucrit Australia, with a review of the Australian 
*peci&s of Brachionus arid Keratella Ay.\t J Mar. 
Freshwat. Res. 30, 237-253. 

(198 1) Zur Morphologic Sysitraalil: und 6kolo^ie 

von neuen moTuwnorucit Raderliercn I Rotatoria) aus 
<lcm 'iberschwemmungsgebtel des Magela Ck in der 
.Alligator River Region* Ausiri*lietis N.T.L Qsnabmcker 
Naturals, Mitt 8, 97-J26. 

<I9W) 1>M5 Radcnier Portral. Triehot/u) felrartii. 

unu vcrwandtc Arren. Mik tokos mos 73. 113-118. 

& Roblrtsuis. B, (1983) laxonomic studies of I he 

Roiifera (Hhylum Asclielminthes) from a Central 
Ama/onian varzea lake, Lago Catnaleao (llha dc 
Marchanlaria, Rio Solimocs, Ainazonas, Bra?ir> 
Amazomotui 7. 225-254. 

A Shiei., R. J. (lygOi New Roiifera from 

Australia Trans, fi .y<jc. X Aust. 104, 133-144, 

<I983) Morphology and ecology of new 

rnonogonont Rotirera from the AHtgator River* region, 
Northern Territory- Ihid. 107, (09-121. 

.0966a) New Roiifera <A*chelminrhes) from 

Tainminia. Ibid. Ilfl. 93-109. 

< 1986b) Roiifera from Australian inland waters 

I Bdelloidea (KolinVra: Digononta). Autt J. Mar. 
Frvshwffier He*. M, 765-792. 

(l9S7ai Tasmai*iian Rotdcra ^fi'iiiities with the 

Australian fauna. Hydrobioioyw 147, 3 1—43. 

(1987h> Roiilcra from Australian inland waters. 

(L F.piphanidae and Bnjchiotiidae (Ronlera 
Monoaoncntta). tnvert. luron. 1. 949-1021 

& Tan L. \V, (1988) New rotifers (Kmifera) irom 

Tasmania Trans. R. Sat S. Aim. 112, Ii9-i3|. 
Ko/ar, L. (1914> Zur KotatorlorttMHia J" 
Tofmoorgfwasser, ^ugt. t. Frgaftmue dieser fauna 
GalJ/tenx AkW. Am,. 4J, 423-^425. 
KnttvCiVA. L. A. (I959J (A siudv of the rotifer fauna o( 
Latvia.) Fauna taiv, &S.R. I. 2II-23L |Rn«iauJ. 

(I97(i> [Rotifer fauna o^ the USSR. Subclass 

HuroraiorJal- Fauna CCCP 104, t .744. (Akari, NauK 
CCCP» Leningrad.) |Ru\siun| 

£ V,\ssat<v\, a. L. 09821 [New and endemu 

rotifer* of (he genera Synchacta and Emit/anis in die 
faunaof BajUIJ pp. 43-5?! in G. I, Gatazii iEd | Nqv« 
o faune Baikal 3 "Nauka" Novosibirsk. (Russian] 
LAt'TraaoKN. R. (1913; Rotatoria- Hattrtwanerkuch d. 

SltourwisZ *. 483-495- 
LciasLwtr, R. (1924; Duv Mannchtn vvn Euchianis- 

dttfiexa. Toot. A/fz. S9, 4ii-5l. 
LcvANork, K. M U894> M^lerulen zur kcnntnis der 
Wa^serl'auua in der Umpefoins von HeUingtors, twit 
beMtnderer Hentcksichu^itng der Mectcsrauna. J1. 
Rotatoria, Acta Soc Fauna."Fhru ftMfia 12, 1—70. 
LtKks, R. (1912) 'Zur Koralonenffluna Wcstpreussens'. 

Manfrlivi. P 0927J Prima nota inrornoally fauna delta 
Goffl dj Bcrtcmica (Adda). Roitferi. Boll, pesxa, 
piscicutt, e UirobwU suppl. tttem. L Ser. Bk, ?-5* 
Mutt Fit. O. V. U7731 'Vermiuro terrtstnnm el 
Nnviaiilium, •teu animal lum Infusiorutn, 
helmiulhicorum et tesweeorum. nort marinorum, 
suconcta Hisiona 'Vol. I. lufuioria. (Copenhagen and 

(1776) 'Zoologiae Danicac prodromus, ?cu 

animal iurn Daniae et Ncrvegiae indigcnaruni 
charaetcres hon^ina el synonyma imprimis popularitim.' 

I M 


Murkav, J. ll>13) Notes on die family CmUypindae «'/ 
R. Miervs. Sue. Set. 2. XV 545-564. 

Mvi rs ( F. J. (1930* The rotifer fauna of Wisconsin. V. 
The Rcncra Euehfairb fiild Monomwata. Trans. Wis. 
Acad Arts Sc£ t*//. 25. 353-413. 

(19,34| I he distribution of Rotifera on Mounl 

Ueseri Island. VII New lestudinellidae of the genus 
Testudinelki and a new species of Bruchumidavof the 
genu* Trichotna. Amer. Mus. A'ov. 761, 1-8. 

(1W») INammoluuval rotifer* of Ijenape and 

Union Lata, New Jersey. ibiO $30, 1-27 
(1938) New .specie* of Rolilcru from the collection 

6f rhe American Museum of Natural tlittory. Ibid, 

LI91, 1-17. 
P.vmsfc. A. (19*63) Observa/ione s-ullv-ieiouonU. 

sperimemale tn Euditanis. Aiti Accad. Nor. Lincet. 

Rend Ci SCI, Ser, K 35, 611^-6 » 5. 
____ (|966] The genus Euchtunis (Routtoti*) in Ihc 

marsh of Fuceeehio (Central llalyfr with description of 

a new speeiev HY'JnsJifologtu 27, 3-28-3 36, 
Pt'iKK, B. <I962) On the taxonomy and woloty of 

oenrhie and neriphviic Rotatoria. Zooi Bidra^. Vppx 

33, 127-422. 
PtKi v M. 0S43) Mifcro^knpiselie Oreamsmen der Alpen 

und der iulier.ischen Schweiz. Mitt. Noturf. Gvs. Bern 

164/165. 153-lVfii 

(18501 Neut Radoiicre dei Sehwen*. ibid. 

Ift9/I70, 17-22. 

(1852) System der tnfusorien. that, 231/232, 57-68. 

K«manf. A. (1929-1933) Rotatoria In "Ktasscn u. 

Oidnunaen des Tterretchs" (Pd. H. O. BrcmtU Vol. 4. 

pp. 1-576. (Leipiif.) 
Kiooi-k. Mi oi (I960) Ecologiich-fawiisiiiche studic van 

dei radCfdieirjes van de Camargue. VerM KonninKL 

vtaamst akud. weienschap. let fef/jj-. 65. 5-2IM- 
11972) Rotatoria. 77te Zttotojiv uf ktland II 13, 

, (l977>RounoiiaoUlieOnl>hc3ii Hegiofl. Studies 

ow the fauna ofCuracoo and viher Catibbeun idond^ 

52, 73-134 

.(1981) Roii fern. Hydrobtoiogieat survey of the 

Lake Htm%\mdu luapula River basin 11, ISl pp, 
Riikoia, H. (1925) (Indev of rotifer* in the bay of Maualu 

and its immediate environment, with description of a 

new species, Maerochaei us esihvni\us.) Vwis, AfljUJ 5g6 

Um'i'. lartutt, 1-12. fFstonianl 
Rom wm n I tl935j Die K.iderrierfauna der Bukuwina. 

Ihtc Syslcmalik B)olot;ie ung seograpnische 

Vcrbreuune. Butt fittv Stiinte Cernauri 8 (1^34). 

(1940) RfclCWC'tanna K urn aniens TV. Zotu\ Ant. 

130. 272 259. 
Knnrxv.M nRunr.vcu. I . |I9N0) Radertierfnuna 

Rumania* VI Ibid. 104,450-68, 
ki'tx.seiu I- <1960> Rotatoria. Fauna Rep. flop A*o/?i 2. 


RUTTNrft KOMVICO. A (197:) Ml Koifltouft {fyft 

Zaoptankton der Btnnen^wasser. Die Binncnaewsjer 

2fK yy-2M. 

(1974i rianktoit Roiifei-is Biology and lasouomy 

^ Suppl. I 1-146. 
Vir.A, fJi 0952) C'nmribuiion al conocimiemo de tos 

RoiiiVros »ie tapaJA I Rondos tie las ^gfi9S 

emitittenlalo Lapaiioles > *u disUibucion. /V/i/. fftSl. 

Hwl. Aphc. BaTcelona II. 
Suin . K I. (lV7n) Associations of bniotno^iiieo Willi 

vveedbed lia.bitais in ;i Inllaboivgol'the Goulburu River. 

Victoria. Attst J. Mar. rreshwui. fibt 27, 533-549 
& K05TL ( W. (197V) KoiifeTa recorJed from 

Australia. Thirty, if *W .S *uu }1fc 57-68 
< 1985) New bpeeiei and new avoid* of Rolifcrtt 

<A.sehetminihes) from Australian waters. Jbid lW^ U|j. 
Sim i K. .],, WAi vi'ft, K. K & Wli I |«MS, W f>. <|y82) 

HlanKton ol \he k^er River Munay, Soulh Australia. J. Mar. Freshwater Res. iJ, JOI-327, 
Skiwima- A- (1903) JNote on truce new ipwileti ui' 

Rotatoria! A*w ftfln, Zoo:. AcudSvi,, St Petersburg 

*. 1^-21. IRussiaol. 

(1914) [On the plankton of the lowvr Voli-«, wiih 

a contribution 10 the QlKSMOti of "l J olatnoplanklou".| 

Tr. tchtyoL lab. Astraehart 3, 3-3J. [Russian]. 
Tait, R. D., Sihel, R. J. tt Kostl. W. (19841 Structinc and 

dvttamics of roopJankmn eonimuuiucs, Altigaior Rfvvij 

reyitm. NT, Australia. Hydrobiokwv 113. 1-13 
Thome. V. C. (ISS7) On certain Rotifera found in ihe- 

ponds of Ihe gardens of the <\eclimaii.sation Socictv, 

Ki.sbana tfw, K SW; Qkl 4» 2K-30. 
(1893) The Rotifera of China. A Raw Mtcrosc, 

$oc„ 12, 145-152 
Vuto-i , M. (1937) -*Ro;atoria, Ole Radenicr/ MKieleuropas". 

- Vols. (Bomtraeger. Stiiiigan ) 
VoRONko^NV. (J9I2) |Ou the plankton of the lakes of 

the Varnal Peninsula. Rotifers and general character of 

the plankton.] Ann. Wu.s. Acad. Se^ Si Prtmbutx Hi, 

1R0-2I4. |Kus*xijriJ_ 

Vkanovsky. M. (1957) Ncikdko virnikov tRotatoiiai / 

ry^ovveli poli mt Sloveu^ku- Biolo^ta Bratislava 12* 

Wam.Ciiia.Ciu. (1961) [^re^hwater Rotifera of China^. 

Inst Freshwater Hydrohii.)!,, Pektnt;, 288 pp. IChiitCMrj. 
\V>a»-M. P. tt898) Faunc rotatorienne du ba^in de Le'man. 

ft*, 5WSV. ZOOi 5, 203-7H5. 
WpsTNtiFKri-l I'NO. ('. (1929) Roratotla Hurujbtteti d 

Zovf I. 1-120. 
Wlsitrk, G. (1894) On Dtstvla tpmifcrt?, ./. Qtuxkctt 

Mier, Ci Set. 2 5, 427-428. 
Whitelegge.T. (1839) List of the inatine and riwUwirtei 

invertebrate fauna of Port Jack.%on and the 

neighbourhood. Proc. R. Soc NSW, 23, 118-325. 
WtCfc/EJSta. A. (18*>3) Rolaloria (wtoiki) Oalieyi Bvtt 

Internal. Acad. Set. Cracmte H892). 402-407 
Wiv.sifcwSKi. J. (1953) Fauna vvrotkoAv PoUki i 

pT/ylegHch. Pot. Arch. Hyrtrobiot, I, 3l7-4Vfi 
Wutti-Hl. K. (1939) Bciiraue iut Kennlni^ der 

Rajderlierftuina Di;uis<;hl;«UCU. IV. Die Raidferttwe dr-r 

Saalc-Elstn Niedeiuug bei .Meriebutt, in okologisch- 

fauni^ri'.cher Deieiliung. Arch. Ilydrobioi 35, 56>-624. 
1 1340) Rotfltoi ten tinigcr ostdeutscher Tbrfmoore. 

An-h. Hytlrobivt. 36, 552-587 

11942) Neue RoUtorienurteu aui deui.%chen 

Mineralqucllen. /.(/id Anz. 137. IS7-2CH). 

(1950 Das Natursctuii^.ebirT nut" dem Tilnrrer 

Selmceberj:. LHe Raderticie des N»tuii\.1iiit^ebielCb 

Arch, riydrohiot. 44, 441-471. 

(1V56) Die Kndcrrietc des ieutels^ct.s net 

rriediiehshauen, Arch Hydo.>bhl, SI, 457-435. 

„ (1960) Die Radertiere sauret dswab^O drr 
Duhener fleidc. 1. Die Rotaiorien des 2adliticmoors and 
des Wildcnhamer Hrnchs. Ibid to. 311-333 

(1961) Die Radetiierc sautcr Ccwaitsei der Dubeiier 
Keide. Ill- Die Re-tatorien des Pres^eler und derr. 
Winkclnuthloi -leiches. fhhl. 5Jt, 72-41)2. 

{ 1 964) V nsei e ^ege hWj | i ge Ke n n ( ni s d er 
RotaloriengaUung Matruchaeius. Ihnnoto^ieu Berlin 
2. 281-W. 

(1965) Uadcrticre aus cinigeu alrikanischcu 

Oewas.sern. Ibid 3 t 347 -366. 
Yamxmhto, K- <l'i53> Preliminary studies on the 
Roiatonan fauna of Korea PtiCij set. 7, |51-ir»4 




Glauertia ohentaiis was described from three male frogs 
collected by W. H. Stalker in 1905 on Alexandria Station 
(19°08' , 136° 43' vide ] ) on the Barkly Tableland, Northern 
Territory, and a female frog collected by G. H. Wilkins 
in 1925 from Groote Eylandt 1 . 

The species was referred to Glauertia Loveridge because 
of its moderately extensive webbing on the toes (Fig. 1) 
and its widely exposed frontoparietal fontanelle. 

Fig. 1. Plantar view of the foot of Uperoleia ohentaiis 
(Holotype, Alexandria Station), 

Glauertia ohentaiis has been reported over a wide range 
in the north of Australia 2 , 3 , 4 . The genus was referred to 
the synonymy of Uperoleia^ when it was shown that the 
generic characters represented extremes in continua 
expressed within Uperoleia. U. ohentaiis was redescribed 
from the holotype and topotypic paratypes 5 but the Groote 
Eylandt paratype was not examined at that time. Its 
conspecificity was called into question because of 
distribution and other data provided in the generic 
revision 5 . No other referrable material was available to 
these authors. 

Later collecting 6 failed to produce further specimens 
of U. ohentaiis at the type locality, and further studies 
on Groote Eylandt resulted in the female paratype being 
referred to U, inundata Tyler, Davies & Martin. 7 

Uperoleia ohentaiis thus remained known only from 
the type locality and from the holotype and two topotypic 
paratypes. 8 

Whilst examining a collection of Uperoleia from the 
Northern Territory Museum (NTM), I found a specimen 
referrable to U. ohentaiis. 

The frog (NTM RA88) was collected by Mike Gillam 
during the Cox River Survey on 2.7.1977 adjacent to the 
Arnold River (15° 43' , 134° 37' ), at night in paperbark 
woodland (Melaleuca spp) in a spring-fed swamp. The 
species was abundant at the locality 9 . 

The specimen is small (probably sub-adult) with a snout- 
vent length of I7.6mm. Toe webbing is moderately 
extensive (characteristic of the species) and there are very 
prominent subarticular and supernumerary palmar 
tubercles on the hands. The head is small in proportion 
to the body and the parotoid glands are prominent. 
Inguinal and coccygeal glands are extensive and coalesce. 
No mid-vertebral stripe is apparent. 

The site of collection represents a range extension of 
about 500 km to the north north-west. 

Parker, H. W. (1940) Novit. Zool. 42, 1-106. 

Barker, J. & Grigg, G. C (1977) "A field guide to 

Australian frogs" (Rigby, Adelaide). 

Cogger, H. G. (1975) "Reptiles and amphibians of 

Australia" (Reed, Svdney). 

Tyler, M. J. (1976) "Frogs" (Collins, Sydney & London), 

Tyler, M. J„ Davies, M. & Martin, A. A. (1981) Aust. J. 

Zool. suppl. Ser. 79, 1-64. 

Tyler, M. J„ Davies M. & Martin, A.A. (1983) Trans. R. 

Soc. S. Aust. 107, 237-242. 

Tyler, M.J., Davies M. & Watson, G. F. (1986) Zool. .1. 

Linn. Soc. 88, 91-101. 

TVIer, M. J, & Davies M. (1986) "Frogs of the Northern 

Territory" (Conservation Commission of the Northern 

Territory, Alice Springs). 

Gillam, M. In Hit, 

MARGARET DAVIES, Department of Zoology, 
University of Adelaide, G.PXXBox 498, Adelaide, 
S. Aust. 5001. 



by Jacqueline M. Nobbs 






There is little information on the occurrence of plant 
parasitic nematodes in arid South Australia. Twenty-tour 
genera of plant nematodes from the area encompassing 
the Northern Flinders Ranges to Cape Jervis have been 
identified together with a relationship between the 
distribution of plant species and the distribution of plant 
nematodes 1 . The distribution of plant species was related 
10 rainfall, i'ollowing on from this work, a survey of the 
arid region (mean annual rainfall less than 250mm) was 
undertaken over a four year period (1983-1986). 

The soil was sampled from under native vegetation, the 
nematodes extracted using a modified Baermann's tunnel 
technique 2 and where possible the nematodes identified 
to species level. At many sties few mature female specimens 
were extracted, so identification to genus only Wats possible 
(Table I). 

There were several genera of fungal feeding lylenchids 
identified which included Aphelenchus avenue, 
Aphelenchotdes sp., Coslenehus c.f, costaius, 
Neoiy/enchus s.l. Tylenchus s.l, and Dhylenchus sp.. These 
are not included with the plant nematodes in Table 1. 
Twelve genera of plant nematodes were identified from 
the arid region as well as juveniles and males of the 
Heteroderidae which were not identifiable because of the 
absence of females or cysts. 

The most widely distributed plant nematode was 
Ti'Ienchorhynchus tobari (Sauer & Annells, 1981). OJ' the 
374 sites sampled, 63'Vo contained T. tobari. All other 
species occurred at less than 8% of the sites. More than 
one ta\on was present in 26°!a of the sites and a similar 
percent of sites had no plant nematodes present. The arid 

regions arc an adverse environment for most organisms. 
To survive, organisms must be drought resistant and also 
respond rapidly to the stimulus oi rainfall. Some 
nematodes are ideally suiled to the arid region as they £ftO 
form an anhydrobiotic state in which metabolic respiration 
is greatly reduced but are quick to revive with addition 
of water 3 and begin to reproduce rapidly after revival. 

77 toban (Sauer & Annells, ]9Hl) is a widely distributed 
nematode within the region and must be well adapted to 
the environment. The species was first described from a 
site near Wentworth, N.S.W, ". The species was cultured 
by planting seeds of certain species of Chenopodiaccae 
(a common family in arid Australia) into soil known to 
contain T tobari. 

Studies of host/parasiic relationships and environmental 
effects on nematode population growth will be presented 
in detail in another paper. 

1 Rtay, F. & Wallace, H. R. (1981) Ncmatologica 27. 

v Kcliindkr, A. E. (1961) Plant Disease Reporter 45, 

747-74 K 
! freckman, I). W., Whitfonl, W. G. & Steinbcr^r. V. 

(I9N7) BioJogv and t-ertilitv of Soils 3, 3-10. 
« Satier. M, K. & Amirtls C *M. (1981) Nematologica. 27, 


IA< QUfcLlNH M NOBBS, CAD] Institute of Parasitology, St Albans, Herts., U.K., AL4 OXU 

1 able 1. Number and percent oJ sites from which plant nematodes (PP) were, extracted in the years 19X3 to I9H6 

Ocnus and Species 


Percent of 
Total Sites 

Tyienchorhynchus, tobari Saner & Annells, 1981 

Pratylenchus sp. 

Morulaimus sp. 

He/icotylenchus sp. 

Scutedo/iemu sp, 

Hadophotus sp. 

lylenchorhynchus vclatus Sauer & Annells, 1981 

fyfenchvrhyrichus sp. 

Jtotylenchus sp r 

nophhiimus sp. 

Morulaimus genkuiatus Sauer, 1966 

PurafonRfdorus sp. 

Heteroderidae males and juveniles 

tyeutedonema iaevi/lexum Phillips, 1971 

Jyfenchorhynchm urmutatus (Cavsidy, 1930) Golden, 1971 

ScuteUonema minuttmt Sher, 1964 

a-Ioiy tench us hasmlatus (Golden. 1960) Siddiqi, 1963 

ffelicotylenchus variabilis Phillips, 1971 

fiadaphohts crenatus Colbran, 1971 

Paratyfenchus sp. 

Hemicychophora sp. 




total number of sites sampled 

Sites with >1 genera of HP 
Sites with PP 
Sites without PP 






VOL. 113, PARTS 3 & 4 
30 NOVEMBER, 1989 

Transactions of the 

Royal Society of South 



Koste, W. & Shiel, R. J. Rotifera from Australian inland waters IV. Colurellidae (Rotifera: 

Monogononta) - - 119 

Short, A. D., Buckley, R. G, & Fotheringham, D, G. Preliminary investigations of beach 

ridge progradation on Eyre Peninsula and Kangaroo Island - - 145 

Pledge, N. S. The occurrence of a diprotodontid marsupial referable to Euowenia grata 

De Vis in western New South Wales ------ 163 

Austin, A. D. & Allen, G. R. Parasitoids of Uraba lugens Walker (Lepidoptera:Noctuidae) 
in South Australia, with description of two new species of 
Braconidae ---------- 169 

Barker, S. Contributions to the taxonomy of Australian Buprestidae (Coleoptera): New 
species of Astraeus and Stigmodera (Castiarina) and a key to Astraeus 
{s.s) ----------- 185 

Jenkins, J. F. & Hasenohr, P. Trilobites and their trails in a Black Shale: Early Cambrian 

of The Fleurieu Peninsula, South Australia ----- 195 

Gowlett-Holmes, K. L. & Holmes, N. J. C Rediscovery of Primovula (R) heleneae Cate 
and description of a new species of Crenavolva from South Australia 
(Mollusca: Gastropoda: Ovulidae) ------ 205 

Davies, M. Developmental biology of the Australopapuan hylid frog Litoria eucnemis 

(Anura: Hylidae) - - - - 215 

Brief Communication: 

Beardsell, G. R. Hybridisation of Litoria chioris and L. xanthomera. (Anura: 

Hylidae) - 221 






VOL. 113, PART 3 



BY W. KOSTE*, & R. 7. SHIELf 


Diagnostic keys are given to genera and species of the rotifer family Colurellidae {Colurella, 
Squatinella, Lepadella, Heterolepadella) recorded from Australian inland waters. All species are 
figured and available distribution data and ecological information given. Some widely-distributed 
taxa not yet recorded from the continent are included. 
KEY WORDS: Rotifera, taxonomy, Australia, known species, Colurellidae 

rrorwrnttrh of the Roy«{ Socwt" of S. Mist., (1989), WMh H9-U.5. 


by W. KOSTfel & R. J. SHtLLT 


KostFj W. ( & Shjki., R. .1, 0989) Rotitcra from Australian inland waters. IV. Colurellidae (KnttfVni: 
Monogononta), Trans. R. Soc. S, Abst, 113(3), U9-143, 30 November, 1989. 

Diagnostic keys are given to genera and species of the rotifer family Cofurellidae {Colarellu, SuuvtiHCldK 
Lepadella, Hetcrolepadclla) recorded from Australian inland waters. AH species are figured and available 
distribution data and ecological information given. Some widely-distributed taxa nor yet recorded trom 
the continent arr included. 

KfeY Womw Rnt.k-ra, i^nnooiy, AusimU, known specie*, Colurellidae 


The monogonont rotiter families considered to 
date in our revision of the Australian Ron fern (Kostc 
AShiel 1986b, 1987b, 1989) have included the moM 
common planktonic taxa in Australian inland 
waters. Here we review the Australian represen- 
tatives of the family Colurellidae, several genera of 
which have widely-distributed species in plankton 
and litloral collections in Australian waters and are 
considered hcieoplankters {sensu Hutchinson 1967) 
rathci than accidental incursions, i.e. 
pseudoplanklon (= tychoplankton of Ruttner- 
Kolisko 1974) (eg Lepadella patella). In general, 
however, xhc Australian representatives of the 
Co I u ret lidae ( Cohirella, Squat irtella, Lepadella, 
Hererolepadelfa) are littoral (epiphytic or 
epibeuthie) in habil. They are encountered 
commonly in the shallow vegetated areas of 
biltahongs and other wetlands, where diverse 
inaemphyte associations effectively partition the 
habitat, providing abundant microniches. 

Diagnostic keys, brief descriptions and figures of 
all known Australian representatives of the family 
art provided lo enable identification to species. 
Some widely distributed taxa not yet recorded from 
Australia are included. The format follows that of 
earlier parts; diehotomous keys are followed by 
individual species' descriptions and known 
Australian records. Relative abundance is indicated 
by: "common" (more than 30 widely dispeised 
records), "uncommon" (10-30 localities), "rare" 
(< 10 records). Brief ecological data are included 
where available. Fatly Atistralian records cited in 
Shiel & Koste 0979) are not repeated here, nor are 
the majority of northern hemisphere citations given 
bv Koste (1978), which contains a detailed treatment 

5. I.udwifc-Bnll-Stra.vse. Quakenbruck D.-4570. Federal 
Kopiihlie of Oermany 

Yltirrav-Darhng treslutaier Research Centre P.O. Box 
9?l. Alhlirv, NSW 2640 

of the Ronfera outside Australia. Keys to families 
are included in Kostc & Shici (19K7b). 

Family Coluielbdae Bartos 

Loricate rotifers, the corona in all genera with 
a membraneous cap or semicircular shield 
(retraetible in all except Souafineltay, no lateral sulci; 
Colarella and Puntcolurella with ventral sulcus; 
irophi maUeate; ventral plate with caudal foot- 
opening or enlarged for basal fool segment; toe* 
slender, sharply pointed. Four genera; Coturella, 
Lepadella, Squarinelfu, l^araeolurella. 

Key to genera of the Family Oilurcllidue 

1. Lorica laterally compressed, with venual or dorsal 

and ventral apertures (Fig. 2) 2 

Lorica dorso-ventrally flattened, without such 
apertures . ] 

2(1). Lorica a single structure, with ventral apcittue 

(fig, J1 M ; , Cohtrella Hory de St Vincem 

I orica two plates, *i|h dorsal and ventral apertures 

(Fig. 2:8) Paracolurelfa Myers (not known 

from Australia). 

3(1). Hood or head* shield large, non-retractibl- 

(FigJ) Squalinella Bory de St Vincent 

Hood small, retraetible (Fig. 5) . . - ,, 
... - - Lepadella Bury dc St Vincent 

Genus* Cohirella Bory de St Vincent 

Cohttvlta Bory de SI Vincent, 1824, v.ZQX 

One piece lorica, in latcial view oval, egg- or 
hatchet-shaped (outline of lorica differs according 
lo degree of eonlraction, particularly ventral 
margin); anterior lorica margin more or less 
I mutated or rounded; posterior margin rounded 
gradually merging into blunt, straight or curved 
projections, in some cases sharply pointed, offset 
whv 'ke extensions of posterior margin (Fig. Ich 

120 W, KOSTE& R, J. SHIFJ 

ventral cuticle between mussel-like lorica borders eeotypie variation ["MiUieufaktoren^fHauer 1924)], 

delicate, membraneous, may be distorted in e.g. the variability in the lorica ends in C. uncinata 

preservative; corona of the Euchhnis type (cf, Koste or C adriattca. Eeotypie variation is common in 

& Shicl 1987b); foot with three or four segments, the Australian brachionids (Koste & ShicI 1987b), 

basal segment mostly pliable and often of indistinct and variants arc known in the five species of 

structure; distal foot segment (before toe) with Colttrella recorded from Australia. It is likely that 

dorsal sensory pore; toes of variable length, species other species occur here; in view of their small size 

with long toes more compressed laterally; toes (most < 100 /mi) eolurellids are easily overlooked 

separate, however may appear fused as frequently in detritus-rich samples. 

are stuck together; gastric glands mononucleate; 

trophi malleate; eyeless or with two eyespots, which Key lo species of Ihe Genus Calureila 

may or may not be red-pigmented; lateral antennae , , 

. , xvt -i ! * i I. Lonea surface smooth .2 

very conspicuous; ma e little known and f - , , w ... . .. . r . . «*!„ 

■*,.„. f J , . . t, ». . ■ i • I.onca surface with loiiEitudinal furrows or tacets 

msuliiciently described. Resting eggs retained in c (esse t a(a (Gla.scoti) 

lorica of parent on death (cf. Fig. 2:3d). ^ 2(1) Loric . a vaJve ^m\) slender, posteriorly with 

Variability in (he described species requires variable apices 3 

investigation, particularly whether the morphology lorica valve rounded, plump 

of different race* ("var.") or forms represents C obtusa (Gosse) 

Fig. I;l, Calureila adriatka (Lhrenbery): (a) lateral (an = anus: au =- eye; ci s- >ubiianeous Vgfi! nix = masiax; 
vi - vuclianum); (b) lateral, contracted; te) ventral; (d) posterior lorica showing loot and toe (D.S. ■« dorsal sctiscsrv 
pit); (fl=-ij lateral view 1 ; showing variation in lorierJ morphology, 2, C, colurus (Ehrenberg): (a) dor.sal: (b) ventral; (e) 
laleial; (d) male, latei;*] ft, ( - olifit\u (Go.ssl-). frfj dorsal; (b) venrrai; (c) lateral: to"! another morph, lateral, la, after 
CordiMlfcM); lb, Ad, after Carlin 11939); le, d, ahci W'ull'crt <W50>; le-t, 2a-e, after » lauer fl l J2f); 2d, after Wivmewskj 
il95-i); 5a-q, after Haider 0924). Scale lines M> />m, 


3<2). Posterior end oMonca With raised lines terminating Colurella (Gosse) 

in vanublc lupenng, downward-curving (occasion- g[Q [ ^ 

ally lightly upward-curving spines; lorica hlrfj 

(length/height ratio <l.8); toes short. Coluntsobiusus Owsc to Hudson & Hoy**, 1886, p. 103, 

■ • . C uncinate (Mullet) Fi*>. 26:3. 

Posterior end of lorica broadly rounded, or blunt- Colurella otetusu after Hofcten. UNA K, 84 
*pmed or tapering, lorica low (ratio >L8); toes long 

4 Diagnosis: Lorica outline plump; margin of head- 

4(31. Lorica valve posteriorly rounded, or at tnost aperture directed obliquely from ventral lorica* 

extended m a blunt apex (Fig. l;2c) V enrtal lines variable; head width appro.v hall loric;! 

. ... c coturus (Lhrcnbcrg) | cngth 0r 5Tna | [cr . evespots with or without 

Lunca vahe posteriorly with curved spines (Fig. 1:1c) pigment ^»* " m » '-' wiwmr 

Length 60 /an; width 2K /<rn; toes 17 fim. 
Distribution: Cosmopolitan between aquatic mac- 

Colurella a<1mtica Ehrenbere r^hytes; panccmunental, uncommon. 8.0-29.O°C, 

F ,i- i.i PH 3.4-7.5, DO 11.2 rng I J, 36.4-1600 uS cm \ 


Colurella adriatica fchrenberg, 1831, H£ J:J Literature: Evans 1951; Koste 1978, 19X1; Shiel & 

Diagnosis Extremely variable lonca size and 

pbenotype, particularly points at posterior end of Colurella tesselata (Glaseott) 

lorica. These are the only features separating C FIG. 2:1 

adriatica from C cu/urus, Pejler (1962) noted 

transitional forms. Different ecotvpes were des- PW**f /**&** fflfcwli, IK93, p. 74. 

cribed by Hauer (1924), with increased lorica size Co!urelUi t*9#t$* After Wflfmp » y < 19 - P. Si 

and icie length correlated with increasing salinity. Diagnosis: Lorica with dorsal keel; pattern 

The measurements below encompass the range of with ribs on lorica surface; toes relatively short 

ecotvpes (Kostc 1978). Ungln 55-64 M m; width 24-31 m\ height 32^41 

Length 85-113 /un; width 29-18 /mi; height 44-66 iU m; toes 18-27 t m\ 

,un; toe length 28-48 M m. DMrihut ion "iCosmapol fan (not recorded I rum 

Distribution: Probably cosmopolitan. Euryhalinc. N.Z,), sporadic m oligotrophy waters, mostly in 

Rare; tresh and slightly saline waters or southeast moss, also in psammon. Single record from Coghills 

S. Aust., Tas. and western Vic. Ber/.ins (1982) Creek, Tourellcn Vic. 

recorded C adriatica as widely distributed in eastern Literature: KomV 1978; licr/jns 1*J&2 
Australia. S.(K30.0°C t pH 5.5-10.5, dissolved 

oxygen (DO) 6,0-11.2 mg I \ conductivity (K. 7r ) Colurella uncinate uncmata (Muller) 

57-13,940 pS cm l turbidity 21-67 nephelometric ''<•• 2:2 

turbidity units (NTU). A1 . , „ „ 

M*5S« Ridder 1972; Kos.e 1978. 2£2* ""'"'""r u '■ ?*& "\ . 

Colure/tu uncinate alter Haoing, I91J. p. 3D, 

Diagnosis: Lorica posterior end Gradually merge* 

Colurella colurus (Ehrenbcrg) into downward-curving short spines, head-opening 

FJO- l«2 smaller than greatest lonca height. 

Momoo colurus Ghrenherg, 1830, p. 44. . Wh 77-106 /t m wktth30,uT,; height to 60 *m; 

Colurella coluru, after H™. £ 3 p . 29. S^ f? */? "^ £* ^V M 

Dtf//7/)w/;o//.- Cosmopolitan, eurvhalinc. Common 

Diagnosis: Distinguished from C adriatica only by pancontinentat. 8.5-24.5°C, pH 4.8-8.0, DO 

the rounded or bhint-spined postetior lorica. 7.8-1 L2 mg I \ 20-1600 //S cm , 3-160 NTU. 

Resting egg and male known (Fig. 1:2J). literature: Koste 1978. 

Length 71-110 ^m; width 25-37 ptir\\ height 39^55 Comments; Two morphologically distinguishablc 

^m; toe length 25-40 /im. tasa described originally as distinct species* bu: 

Distribution: Cosmopolitan, widely tolerant, ooau- regarded by Koste (1978) as rt torm" variants o\' C. 

ting in fresh, arhallasie saline and coastal wateis. uncinata also are known from Australia. 
European temperature 1 .0-28,0°C and pM 4.9-J0.0. ^ w „ , 2 • j 

Ridder (1972) records C 2mi ftom 15.035 J CI ^^ ^^^(^ (Ehxenberg^ 

i - Rare; Tits.. Vic. U.8~16.0°C ( PH 5.7-M. 

415-2130 /tK etn '. Colun^ bicuapidmus F.hrenberg. 1832, p 129. 

Liieraturt: Koste 197$, Colurella unrumtu f. hiempidata after Moche, 1^39, rx 





Diagnosis: Lorica width up to twice thai of 
nominate species; highest extension ot lorica al 
head-opening; lorica fcridi short, most curve 
downwards; toes relatively short; resting egg with 
punetiform cavities 

length 60-108 p, width 53-62 ^m; height to 
60 /mr. toes 111—27 ^m. 

Disinflation: Cosmopolitan in littoral of fresh and 
athollasic saline waters, also marine littoral. 
Uncommon: N.SAV,, Qld, S. Ausc, Tas.. Vie. 
IO.O-23.5°C f pH 4.5-7.5, 53-927 j$ cm \ 
Lhenjtun': Kostc 1978; Sluel $c Ko&tc 1979; Green 
I9KI; Begins 1982. 

CohtreUa unnnata deftexa (fchrenhenM 
FfG. 2:4 

Catitrus deflesux Ehtcnbctg, 1834, p. 203. 

Ctsiurelfo uminoto I. dtflv.xu after Meucbe, 1919, p. 400, 

Diagnosis: Lorica t:nd robust, elongated, turned 

outward, mosl directed venlrally. Very vatiable 

Length 60-100 jjrth; width to 55 ^m; height 35-55 

iim; iocs 17-25 /*ni 

Distribution: Cosmopolitan, sporadic, enrvnaline. 

Rare; Qld, las. Bjy^KflFC, pH 6.5. 

Literature: Shici Si. Koste 1979; KoueA Shiel 1957a. 

Not recorded from Australia" 

C. dvnticuudu Carlin, 1939. Scandinavia 

C. dicerura (Gosse, I8&7). M Eumpe, murine 

C h'ttidtnbar^t (Sieincelce, 19]7)(h'U. 2:6). Probably 

C. %vopiutu Donncr, 1951. . KurojHr 
C warlttovi Althaus. 1957. Black Sea, mesopxammal 
C mwidodiKtvlos Alihaus, 1957. Black Sea 
C mucranidatQ Ahlstrom, 1938, U.S.A. 
C. ob/oHiw Doutier. 194.3. Europe 
("'. oxynntda Carlin, 1939. Scandinavia 
C paludiua Carlin, 1939 US.S K. 
C Audrta Abhaus. 1957 (Fig. 2:7). Europe, N./. 
C\ sitthtra Carlin, 1939, F.urope 
C. stdaitti fSienrno*. l898)(Fiu. 2:5). Probably 

cosmopolitan, warm lilenotherm, acidophil. 
C, urucauda Lriksen. 1968. N. Europe, 

Genus Squal'tnella Bory de St Vincent 

SquathwlUt Bory de .Si Vincent, 1826, p. 87. 

Body cylindrical to ovoid; head and corona under 
hyaline semicircular shield {fig. 3:1), which is 
pointed apieally in some species, also with 

oiangular side plates ("ears**); dorsal lorica smooth, 
also with occasional symmetric longitudinal libs: 
ventral lorica smooth, with aitached shield-like 
plate, with one or two scales, or also with keel; 
posterior lorica rounded, truncated or tapered to 
variable point*; some species with long donsai spine; 
short spine in median line of posterior lorica may 
be present; foot 2- or 3- segmented with long 
pointed toes; small thin spine may be present at have 
of toes (Fig. 3:1c): symmetric matlcatc trophi (F'ic. 
3:6); gut without peculiarities, dorsal and ventral 
antennae small; two widely separated distinct 
eyespots with crystalline bodies. Male and resting 
egg not yet described. Ecology of Squarinelta little 
known. Koste iJ978;174> notes genus restricted to 
littoral, with sortie spp. acidophils. Four species 
recorded from Australia. For details of other species 
and variants, see Koste (1978:175-179), 

Ke> to species of the genus Squatiiteiln 

I With dw*al spine. .. .2 

Without dorsal >pme .. ,,, , ..) 

2(1), Foot 1 -segmented; short spine ai hase ol toes 

& tontiisptnutu (Talcm t 

Boot 2-segmcntea; spine absent 

5. leydtyii (/-achanaVi 

^{0 VV'iUi a short spine above the base of the lots 

. S. rostrum (Scbmardaj »pine above toe bases 

V. intuitu (KhrenbctfU 

Squatineda lortyispinata (Tat em) 
FIG. 3:1 

Swphatmps irmnkpimmis talcrn, 18<\7, p. 252, PI 10, i"\^ 


Squaiinvlta ionpjsphwtQ alter Harumj, 1913, p. 96, 

Diagnosis: Characteristic tong, thin and curved 
dorsal spine. Pejler (1962) noted two short spines 
flanking dorsal spine (Fig.3;3a). Ventral side 
sometimes wrinkled. Ratio dorsaJ spirte:body length 
0.8. Foot three-segmented. 

length R2-160/un; dorsal spine 96-1 16 ^m; toes 
MO mi spinc(s) at the base of the roes 4-5 /*m. 
Distribution: Widespread but rare, mostly in moor 
waters between Sphagnum (Koste 1978), We have 
not encountered this species in > 2000 collections, 
however it was recorded by Whitelegge (1889) from 
N.S.W., also by Evans U951) from Victorian waters. 
I iterative; Pejler 1962; Koste 1978. 

He. 2-1, ( 'ofuretla tessetwa (OWOH): (a) dorsal; (b. c> lateral. 2, C ttttamiia utKniaiu (Mullen (a) Uuerat; tb) doi.^aL 
tct male, ventral; fd) male, lateral. 3, C utuitwta hicusptdtuu iFtnenherui: do lateral; <b) utxjthcr morph, lateral. 
(O '.eittriil; (d) wniral, with resting egg. 4, C. utKhmw deftp.xo (Hirenbeic): (a) lateral; (b) hubii, dorsal. 5, C suhutu 
(Sfcumtw), lateral. &, C hindeniuir^ SieinL-cke- (a) dqrsat; (b) lateral. 7, C. sattiw Alihaus (aj lateral: (b) \cnhal. 
M, MtfWI idurciiit /nxutttf Mvcis: (j) lateral; |h) ventral. In, b, 2b, 4b, after Hauer (1924); Ift ^a, 3a-t, 4a, 5, .il'lei Carlin 
(mn 2e, U. .Hl'iei WVber (I898»; 3d, «a. b, atier Koste |I9"«); 6a, b, alter Siemceke (1917); '/a. h, after Alliums (1957* 
Seale lines 50 ^m. 


W. KOSTE <fc R. J. SHIEL. 



Squarmdia teydlgii lon&isetu (Ponrriotj 

v\c\ V4 

Squai'meda tcydiyu f. tonwwa Pom riot, 1971, pp. 99-100. 
lie R 

Diagnosis: Robust body; dorsal spine longer than 
body; two foot segments; no spine on distal end of 
terminal segment (at base of toes). Much larger 
animal lhan S. longispinato. 

Body length 210-235 um; dorsal .spme 270-37S 
um: ratio spine:body length 1.6; toes to 37 ^m. 
Distribution: Rare in submerged Sphagnum. 
MyriophyUum, etc. Single record I rum a pool near 
L. Garcia, weMern las. I7.0°C\ pH 3.L 80.6 ■ ptS 
cm ', 

Comment; The nominate S. teydigii is not recorded 
from Australia, It is distinguished from .V. 
lon^ispmuta by loot morphology and from the ssp, 
tonpjseta by its markedly smaller dimensions (body 
length 110-110 ,uu; dorsal spine 90-110 /mi; 
spine:body ratio 0.8-0.95; toes iOamj. 
tJwwure: Koste 1978. 

Resembles S. rostrum* but lacks spine at base ol 
roes. Hendshield semicircular. 

Body length 134-190 ^m; toes 20-30 /*in. 
Distribution: Ponds and pools, often abundanl in 
Sphagnum pH 4.8-6.2. Rare, Old, N.S.W., Vic. 
/ itewture: Evans 1951; Koste 1978. 

Squatinclia rostrum (Schmarda) 
Fife 3:1 

tt\t Hon rostrum Sehmarda. IK4f>, p 20, 1*1. 2. Fig. I. 

Squatinetia rostrum after Carlin-NiUon. 1939, p. 3 

Diagnosis; Caudal lor tea with three long or sho>r 
pointed processes which arise from narrow bases. 
Lateral antennae sunken in dorsal lorica margins. 
Length 150-217 /im; toes 24-28 /mi; headshicld 
to 76 urn; caudal processes to 36 p,m; spine at base 
of toes to 23 ;*m. 

Distribution: In Aulwuch* (i.e. periphyton attached 
to subineryeut stems). f\vo records only, Brisbane. 
Old and from Creswick Ck. Clttncs, Vic. 
Literuture: Thorpe 1887; Koste 1978; Ber/ins |982. 

Sifum'mella mitinv (Ehicnbern) 
HG. 3:2 

Sh'phantif)\ tnufhus fchtertUUA. 18.32. p. US. 
Squtiiineliu mutka after H&mn& IMI3, p 97. 

Pwnnosis; Cauda] mar±yn uf dotsal lot rca semi- 
circular, tongue-shaped, three-lobed. with three 
short spines or truncate (Fig. 3:2d-g). All forms may 
occur in same population. Fool three-segmented. 
Body length 100-226 fim; headshield width 60 
//m; toes to 30 pftL 

Distribution: Furope, N.Z. Rare, from billahongs, 
ponds or dams: N.SM, NT., Tas. ( Vic. I6.5-24.5°C, 
pH 3.8-5.3, 45.5-46.2 ,*S cm r . 
Literature: Russell 1961; Koste 1978; Bcmns 1982. 
Koste & Shiel 1987. 

^(prntmeUit muricu tridentatu (rreteiiiiul 

1-Ri, 3:5 

Srcphanops to t kmtmt\ Fresenius, 1858, p. 218, PL Ift Rg 


Sfiuittinefia muticu tritfrmata after Wis/niewski. 1053, p. 


Diagnosis; Posterior lorica oval, terminating iu three 
pi-nnted, elongated spines of variable length. 

Not recorded from Australia: 

5. hifun-a (Bolton, 1884) in Hrukon & Cosse <lM'> 
liurope, lecland- 

S. mivrodttctyttt (Murray. 1906). Europe, Newfoundland 
in Sphagnum. 

Genus Lepadellu Bory de St Vincent 

t.rpadeila Bory de St Vineem 182<\ f> $vi 

Ootline oval, pear-shaped or circular, more or less 
dorso-ventrally llaitcned; dorsal and ventral lorica 
plates firmly joined; deep notch for head and loot; 
occipital margin of lorica often with granulated 
collar; head with cap-like plate. Dorsal lorica 
domed, smooth, with or without keel development 
also with several ledges or fillets;; lorica margins in 
some species drawn out into winglike extensions. 
Head and foot apertures of different shape; 
differences useful in taxonomy of group. Foot ol 
three flexible segments, one inflexible basal 
segment. Toes short and pointed or long and needle- 
like, similar or dissimilar in length, inserted laterally 
or dorsal-ventrally, in some species completely or 
partly fused. Corona single ciliary wreath with 
lateral ciliary tufts and buccal field. Two lateral eye 
spots, occasionally with separate pigment granules 

Fig J.I. Suuuttm ■//</ wtram (Schmarda). (a) headshield, dorsal; <b> headshield, ventral; \c) dorsal; (d) lateral amenna 
(et vtm.Hl: (I lateral. 2, S. mutuv (Hhrenberg): (a) ventral; <l» laicnll; (c» dor,.al; W-g) variations in cand.V 
i> ( Orpholoy>. 3, S, iottgtspvwtu | latcm); (a) lateral, (b) dorsal; (c) foot, lateral. 4, S. feiitoft Uviweta Pnurrim 
S x. amtfUi tndemmo (Presenilis): CO doiml; (b) lateral. Ih. b, ;d'ier Weho <U898); lc-'t, 2a, h 4a after wulfcrt 
!' n VnfvJ 4 '?' S?*iWBfc 3a, pflfl Pejler (I9«>; 3b. utter Slenroos (1898), 3l. after Myr.s tfatti; |h, Aftd 
WimtA IIM4KK 5, alicr Pouimui (1971). Scale lines 50 ,/m 

\26 W. KOSTE A. R. I. SHI hi 

ncurbs. Dorsal and lateral antennae present, laud 5 < 4 >- Vt - vmral Wf* wirh rvv0 terocfl) s(Lfn$ directed 

general lv dorsaily situated at level of upper border kn W*& ,ue! > M*W {% &£>• : 

Of fuui-opejiinji. Last foot cement usually with ;/"" Vr'-" - '■ ; *' 'fl?. Xcwte* ShUSI 

. , , -r . - it *~ '*t fi A Ventral lorica Spineless: Joes curved 

caudal antenna. Iron n malleate, uncus with rive , . , . u 

. . . * .. . , . . .... — — /,. nen&isst Her/ins 

teeth. Gut with large^ sometimes iobed 6(U Occipital macHiuol head apcMmcptotrudiuit (Fig. 

glanUv Vitellarium larpe with eight nuelcn. 5^ m 

Proionephridia with distinct terminal cells and large Oectpiml margin ol head apeiiurc concavety 

bladder, I iitle known rif reproductive cycle. Some notched (Fig. 5:4a) 7 

males (^0-120 ^in) are known. Resting e^s, wuh 7(6). lorica outline elongate oval to circular (Fig 5;7a) 

prkkiv outer shell, retained in lorica of parent on # 

tl^ath. l.orica outline rhombotdal (fijj, 5:4a), , 

Jepo^elh species are benthic, common in M , " ,~* Ll pto&RAI Kostc 

1 irLdT^JT,* s.^i ,^.,.1- C* **A A *U.*»ii.j m ii.^j „, W ■' )■ Posicuoi margin ot Joiica rouuded or truncate, 

billabooj>s and pond., but occasionally collected a* |wver poimtd L (ri SOa) 4 

incursion species in the planklon of reservoirs and Posterior margin ot lorica 'taper*" 10 « pointed 

rivers, e.g. Z. j^/eVto {Strie! cv *//. 1982) Further -projection extending beyond base bfrac*(>{g. 9:7) 

details are given by Koste (i?78), who notes that , / withatnti Kosre & Shlcl 

all species can be identified I'rorn preserved material, 4<8). Head aperture ^eutrnl sinus witfi parallel cuticular 

although caution must be taken with preservation ribs; dorsaily with a wide fcranttlared collar 

artelacls, patticulaily in Ihe case of lonnalin- — '- tf/flwfer Wttifferi 

preserved specimens' Several of the 45 taxu ot Head Rpwiw* without rib.s or dorsal collar 

tenmwtlu idem died from Australia probably arc tt% ^ L -;-,• * ' ' ' *" ", "',' W^ 9 <M,lHer) 

• -j .•.-• -ii* .u- *fi_ -I ^ (ft)- Ventral Oflca with loneitudma pleats 

rrmideunhed tor this reason. I hese are considered ' Wfl r *T? .« 

,. . . 4 , . . l c - . . r , , A. tnhu Myers 

tn the text. Nevertheless, about hail of the known Vemral lofica unp | eattfcJ ,, appear to be indigenous; Lhegen.iMs thus nO0) Crosvscction with rounded head-like lateral 

second only to Braduotuta (Kosie & Shtel 1987b) matins /.. t&rtjtmo CSienrnrti) 

in ns complement ol endemic laxa. Cross-seeiiou idien-il margins produced to acute 

angled tips \1 

I hiee subgenera are recognized on the basis of toe 12(10. Foot-opening excised dorsath (3 

morphology; Foot-openiug not excised dorsaily 10 

1 3 < 1 2 ) . Extremely wide toot-opening, longer than wide; 

III t-f-pofJcllii (toes ot equal length, entirely corners wuh outwurdly-eiuved point* 14 

separated); l* oul opening wider than lony. no points on 

(ii) Heterolepadella ttoefi of unequal length); corncrs L to, / tf '""" ' ^f'«t 

(iii) Xenoti'putteilu {Cries complelely or partly I'usect). I4(n> - HcAd »W^ ^enlralty with lateral hitu.eotc 

pointed extensions extending beyond dorsal 

For convvnicntc, iho Australian T^" Y , ^ "'"""" <Kos, fj 

, .11 , ■ ,. r 11 rs, ° BWflU points on head aperture.,,, S 

.subgenera are consuleret separatciv ui he to llDWinfi ,«», ,_, , ... , , 

& 'W^wv^ v i^vyi.ih ,^, 4 j Head apeiltuc with convex dorsal margin; last 

K<r ^ loot segment < '/,■ toe length 

key Id species of file genus Upudcth (>. *rr.> < tatuswus (HU^eodorD 

Mead aperture with concave dotsal tnargni; last 

1, Oorsn) lorica without keel or ribs (m^v he lightly loot segment > ',: toe length 

puneiMte, hut otherwise uiioniamemed) ... .1 L. vitrea Shephard 

Dorsal lotica witlt keel aud-'oi lonuiitidmal libs 1ft(t2). Head aperture eticular L. ups'ula Harring 

or oiher urnamentaiion 17 I lead aperture with deep vemral virtus, 

2(1). J .orica length >H lorica width T almott cylindrical J. rotimhuny (I neks) 

{Kifci 4:3) ■ ./.. fW&l&ra Kcrzins tHD- I.onca outline trapezoidal; loiiira uiiiaineiiled 

lorica length <3x lot tea width, ovoid. chcuUUi (:*ppe;n- punctate 01 yranulatcd under 1.1') 

rrapc/oidal or rhornhoidal 3 tK 

>l2'l. UoisalloiicaeioSb-icetton I/? oteircleto highly- lorica onttinc cireuhxt or oval 19 

domed hemisphere {t'\}L 5:2d). ... 6 18(17.) Lsleial extensions ("horns") from posterior lorica; 

OoimiI lonea a shallow dome, or dorsovemmlly rows of tiny rodlike structures covet lurtvM (hiy, 

fladencd . . . : : 4 6:4a| --/. utirtorui Koitc 

4(3). Lotica oval to circular; head ApCtJUtC nolehed t -i(crnl hom\ absent; punciilotm ornauientation 

both dorsaily and venlrally (Fiy. 6;5al of dorsal and vemral lorica (lig. 9:sa) , . . - 

/.. ovuli6 (Muller) , , ./ , tyieti Koste &. Slnel 

Lotica 2x widei in posteiiof thud than anterior;, IU(I7). lorica ourtinc ciictdat; head aperture circular, 
dorsal margin of head aperture straight, venlral veuljally placed; lorica cross-section an inverted 
margin only weakly \ -shaped Umk. saucer (ft«. ft;2). , , . ,< L, linduui Koste 
- * I orioii onrlnir ovotd, head aperture not circular; 



3c 3d 3a 


2a ^b 

Fig. 4:1, Lepadella acuminata (Lhrenberg): (a) dorsal; (b) caudal morphology, ventral; (c) dorsal; (d) caudal morphology 
ventral: le) ventral; |T) lateral; (g) cross-section; (h) ventral; ti) L, acuminata, sexco.stata, dorsal; [\\ L. chorea, ventral; 
(k) L. chorea laieral. 2. L. amphitropis. Hairing: (a) ventral; (b) cross-section; (c) dorsal; (d) lateral. 3 : L, ansusta 
Ber/ins: (a) ventral; (b) dorsal; (c) toot, disial segment, and toes, dorsal; (d) toes, lateral; (e) eross-MXliou; (0 anterior, 
lateral. Ui-d, I}, after Koslc & Shicl (1980); la, f, g, i. k, after Berzins (1982); 2, after Harrins (1916): 3. after Ber/ins 
(1960). Scale lines 50 ,iu\. 

128 W. KOSTii ik 9 »■ SfilfJ 

lorica cross section lightly domed, libbeil or to form an elongated ridge to the end of the loriea. 

LrianuuJur . 20 L acuminata septemcosiata Berztns probably is an 

20<)9). Posterior lorica tapers to an acW p.;.in|, which ecotypic variant. /,. chorea Ber/ins (Tig. 4:Jj, k) is 

may be ridged dorsally %... .. inadequately fieured, and appears to be a 

u • • : ' . L^T^ < LhrenbcT ? ' preservation artefact of L. acuminata. 

!i™^ Literature: Burlos 1955; Russell 1961; Kmte 1978; 

2t ( 20i. Diyidii^ Shield Koste 1970; Green 1981; Bcrzins l!>S2. Kostc 

Dorsal lorica without ftmfoui keel. .22 er at, 1983. 

22(21). Dotwl lories with open-squared pattern (three 

rows); occipital margin straight. . Lepadella amphittopis Harnng 

/.. decora Uemns KIG. 4:2 

Dorsal loi ka unpuitemed, with J-4 jhort ribs iwcr 

root-opening . . 23 / . amphitropis 1 Lining, 1916, jv 543 

23i22r M short rihsat posterior end over toot-opening; L. amphitropts vietotiensis Herons. 1982, p. 10, Fig. 25 

cross-*ee0oti rhombic - . • 

I it mpfiirto/,m Hnrring Diagnosis: Lotiea rhombic due lo 

4 parallel short Tibs over root-opening; cross- curvature of the ventral lorica; lories end with small 

section a shallow dome with lateral concavities notch: dorsal lorica end with one to ihtee shori ribs. 

of ventral margin (Bg. 8:2C) Length 69-76 /an; widlh 54- 56/an; height to 29 

/, quadricarmata (Slcnroos) ^ rn; i ot »^ 20-24 «m. 

24(21) Median keel of dorsal lonca »,rh«ui .side f$S Dismbut ion: In Sphagnum, middle fcuxopc and Nth 

ka'a- ""i+'l" iii Vs'^V-'^Vi'u'rihe " America. Rare; N.S.W., Tas.. Vic. ».5-I7.0*C\ pH 

Median keel with 2-3 pairs of sine ribs , . . 0rt no ' ► 

t wrt/ff (LUCkS) f£*? **£**££<* *,„ „ 

25124, NM.mi keel low and wide, bordered by lateral J Mature: Haner 1958; Komc 1962: Koste & Shiel 

grooves; yranuhiicd >oll;u 1987a. 

( . Z.. rhvtnaoUky (Co&e) Comment: A populat ion from the Tarwin R ,, Vic. 

Median keel lugh, triangular, arising from narrow designated Lepadella amphittopis victoriensts by 

base; no g r» mi Imeil collar. Some forms have fine Bcrzins (1982) on the basis of the deep sinus on the 

.Mnned/pu-.tululcd loiiea surface. ... ventral lorica doe* not differ in this feature from 

• - • - L WWW <P"*nh^« lice nominate species. The measurements of the s.sp. 

NH. Some Huropean forms of L tripteru ate ..*« by ^fiTO W2S ^T'' 'T* R & 
Kostc(l978)«obe.l l ou l l,ou..lO. ( lvovf 1 Mo t nv, a rcknovvn *«^ S^ ^TOW" ~° ^ S* 1- S? Pl 
from Australia, depth 8—12 /<m. ventral sinus depth !4 ^m, fool- 

opening I8-20\I3-I6;im, ioe length 15-18 /an. We 
Lepadella acuminata (rhrennerg) retain subspeeifie status, lot this (axon until the 

... f . .. extent of ecotypic variation can he determined. 

SWopidta acumuiata Lhwnberg, 183-4, p. 210. 
Lepadelht acuminata after l>jjat'dui, IR41, p. 6jL 
IrpadcUa chorea Beuins, 1982. p. 18, Fig. 3r> 

Diagnosis: Dorsal lorica with (wo lateral keels; 
caudal dot sal lorica variable, with small notch or 
strong elongated spikes, which may show short 
sharp or blunt ridge dorsally. Nominate sp. with 
points of dorsal lorica slightly offset laterally. 

Ungth 64-110 juti; width 42-72 /<m; toe length 
IS- 15 ;/.rn. 

Dtstrthution: Cosmopolitan between water plants, 
in liitoral of fresh standing and tunning waters, 
tolerates slight salinity increases, however prefers 
acid waters (Koste 1978). Uncommon; pancon- 
tinemal, IO.()-29.8°C, pH 4.3-7.8, DO 6.1 jug 1 \ 
28-IU20 jtS cm ',8 NTU t Alk, 1,9 rng l*«. 
Comments: Several valiant forms arc known, On 
preset 1 1 evidence these are not considered of specific 
status. In L. ottHW&Ulv wxcostuta Barios (fiy. 4flt] 
ihe dutsal loiiea has six ribs, which run medially 

Lepadella angusta lieryins 
FIG. 4:3 

L. annum Berzim I960, pp. 85-86, lig* 7-12 

Diagnosis: lorica elongated > vcntraily flat, dot sally 
convex; occipital mar,cia with deep U-shapc\l dorsal 
sjaui, deeper V-shaped vcniral sinus; dorsal lorica 
sparsely pntteiate; dorsal pores unusually far 
forward, in anterior '^rd of dorsal lorica; fool 
groove broad, distally not reaching rounded 
pusiennr margin of lorica; foot long, with distal 
segment longer than two proximal segments 
combined; lerminal segment with two small dorsal 
projections (Fig. 4;3c. d); toes long, thin, acute, 
wider ptoximally. 

Length 115 yun (lorica 105 ^m), width 46 /an, 
dorsal sinus 34x18 /on, ventral sinus 34\23 ^rn, foot 
groove 25x18 /an, foot and toes 65 /an (terminal 
segment 16 /an, toe.s 41 ^m), 
Distribution: Apparently endemic. Collected in 



summer from the Plenty R., Morang., Vic. No other 
locality known. Ecology unknown. Supeihciatly 
resembles L. elliptica (Fig, 5:7). 

Lepudella apsida Harring 
FIG. 5;1 

/.. xp.tida Harring 1916, p. 536, Fig, 89:1-3. 

Diagnosis; Lorica almost circular; head aperture 
lound, with posterior border drawn downward; iocs 
very .-.hort, 

Length 70 //m, width 60 /an; toes 12 pfe 
Distribution: In riparian vegetation. N. America. 
Asia Rare; billabongs (NT.), I . Euramoo (crater 
lake) rQld). 28.5°C, pH 5.4, DO 6.2 rng I ' ( 23 n§ 
Literature; Green 1981; Koste 19H1. 

Lepudella benjamini Harring 
TIC, 5;2 

L benjamini Marring 19W, pp. 548-549, Fig. 93:1-8. 

Diagnosis: Lorica outline broadly oval, head 
aperture more or less tubular, narrow, with 
ventrally-direeted margins; lateral lorica margins in 
cross-section willi bulging, beaded rim, or 
downturncd; foot aperture wider than long: iocs 
long, gently curved vcntrally. 

Length 112-114 j*m; width 85-100 ftitk) height 
44-52 nm\ head aperture 32-40 /mi; Font aperture 
24x30 /an; toes 45-48 //ra, 

Distribution: N. America, S. America (f. bruzdienus 
Koste 1972). Rate: I . Boon, Vjti, NT. billabongs. 
20.(K24 ( 5 o C, pH 6.3-7.4, DO 5.1-8.8 me I ', 
■42-1500 u$> cm '. 
I iterature; Koste 1981 

Lepudella ehenxutathi Ko.sie & Shi el 
FIG. 5:4 

L chfnxaiatlu Koste & Shiel, 1980, p. 139, Ftg> 7 u-d, 

Diagnosis: Rhomboidal lorica with rounded 
corners, widest in first third; head opening small, 
not deflected vcntrally; foot-opening semicircular 
at widest point of caudal boundary tff dorsal lorica; 
dorsal lorica weakly arched towards interior; three 
flexible foot-articulations, distalmosl slender and 
elongated; toes symmetrical, lone, needle-like, 
Straight (Fig. 5;4b); lorica cross-section shows highly 
vaulted dorsal lorica with slender downward-drawn 
lateral wings; ventral plate with delicate double 
convex keel (Fig. 5:4d). 

Length 135-140 j*ai (lorica 96 jan); width 92 /an; 
height 4S inn; head-opening width 28 /<m; ftjfcjl- 
opcnine,2Kx2X/im; distal loot segment 19-20 /an; 
Iocs 33-36 fim. 

Distribution: Endemic, only known locality L. 
Boon, Vic. 23.5°C\ pH 7.9, DO 6,0 ing 1 ', 420 & 

cm ' , 67 NTU 

Lepudella comma nom. no v. 
FIG. 5:3 

Lepudella lauisinus f» mueromitu Koste, 1981. p. 119. Hjs. 
Ua, h 

non /.. rmwronukt Schmania, J8.*>, p. 57, PI 13, Fig. 20. 
non Mctopirba muentmuu Daday, 1908, p. 30. 

Diagnosis; General posterior outline of lorica 
resembles /.. tafusinus, with extemely wide foot- 
opentug; in /,, cornutu opening wider in proportion 
to lorica width, i.e. body slightly more elongate; 
caudal margin lightly concave dorsally, ventral];, 
with v-shaped sinus flanked by distinctive pointed 
lateral projections extending beyond dorsal margin; 
points are bifurcate in lateral view (Fig. 5:3 inset}, 
with upper longer tips re flexed vcntrally over shorter 

1 orica length 110 /*m: width 62 /an; loot -opening 
width 40 /tm: toe lengih 32 /on. 
Distribution: Endemic, only record from Nalteen 
Rillabong, Magcta Creek, NT. 29.9*C, pH 5.47. DO 
5,45 trig t \ 44 p£ em J , alkal. 2.A mg I '. 

Lepudella dactyiiseta (Steniuos) 
FIG. r5 

Meiopidkt daclvliseta Stenroos, 1898. p. 165. fife, 3:1. 
l.i'padella rlachiixetu aftCJ Harring, 1916, p. 547, PlM 

Diagnosis: Caudal margin of dorsal lonca flat, even 
or convex; ventral margin with variable sinus (cf. 
L. luiasinus); lorica cross-section always shows vers 
pronounced dome; toes relatively short. 

Leugth 93-100 um; width 60-66 m\ height 40 
/<ny, toes 24-32 /an. 

Distitbutiun: Cosmopolitan in algal-rich standing 
waters, also lake Ihorals. Rare, Vie. 8.5-27.0°C, pfl 
7.0-7.6, DO 7.0-11.2 mg I \ to 575 p% cm '. 
21-52 NTU. 
Literature: Koste A Shiel 1980; Berlins 19S2. 

Lepadetla decora Bemns 
FIC7. 5:6 

L deivru Uerzins 1982> p. JJ, Figs 2a. h 

Diagnosis: Occipital margins nearly straight; dorsal 
lonca with open-squared pattern, arranged In three 
rows; robust pores on last third of dorsal lorica; 
ventral lorica slight; foot-opening wider than long. 
occupies whole postcrioi margin.; iocs straight, 
Length 70 /an; widih 52-54 ,<m: head-opening 35 





;<m; fooc-opening 12x30-32 fi m: toes 24-26 #m. 
Distribution: Only ttTiqwn locality Loddon R„ 
Victoria. Leology unknown. 
Utcratimv Her/ins I9S2- 

Comment: Inadequately figured in Kemns (1982V 
We have reproduced Uer/.ms he^ire, which does not 
show- ihe patterning or pores described above. We 
have not encountered the species however on the 
hum of the above description regard it as valid. 

1 epadellu ellipiica Wulfen 
FIG 5:7 

i rihinwu Wutrbn l93s\ p. 60V, \%, 2L 

Diagnosis; Elongated lorica, hemispherical in cross- 
action (ef. L. angusta); head aperture with shallow 
U-shaped dorsal sinus, deep V-shaped ventral sinus, 
laner with accompanying parallel-bordered lateral 
cutieular pleats; distal foot-segment conspicuously 
long, tapering slightly to base of toes; toes relatively 
long, curved vcntrally. 

length 10N-H5 ,mr. width 70 /.on; distal fonr- 
segment 28 jutti; iocs 20-22 j<m. 
Distribution: Sporadic in littoral decomposition 
zone Known only from a billabong ot l he Magela 
C'k. Jabiluka, NT. 25.5°C. pH 6.2, DO 2.9 mg 1 ', 
62 pS cm '. 
Literature: kostc \9X\. 

iepadelia latusirtus illilgendorf) 
FIG. 5;8 

Mctopaiio so/iJta hlusmus It.lgenoW. IHW, p. 131 H|, 


LcpadcUu iatusmtts alter Hnrnne, 1913. p, ti& 

Dtogttosis; Lorica broadest in middle, tapers slightly 
rti eauh end; erovs-secuon vaulted hemisphere; 
occipital maruiii cowl-like, prniruding; head 
aperture relative^ small; extremely wide fuot- 
opentng, small spikes at matins houked dorsally 

Lorica length HO-92 /tm; width to 64 /tm; hejgln 
35 i» in; head apenure width 24 ^m; toes 24-2K ftm. 
Distribution: In subtropical and tropical shallow 
waiers (bin lecoided from Mt Cook, N.Z.), warm 
sieninherm, Rare: N,L 24.0-27.0°C\ pH -4.5-6.3. 
DO 5 I mg I ', 42 ^S cm '. 
Comment: A variant described by Kosie (1980 from 
Nankeen Billabong, NX, as L, lutusinusd. visenda 
(Fig. 6:1a) had features of L latusinus and L. 

visenda Myers (Fig. 6:1b). Only one individual was 

encountered, insufficient material for furthet 


Literature: Myers 1934; Russell 1945. Kosie 197S. 

Lepadella lindaui Kosre 
FIG. 6:2 

L. iindaui Koste 1981. pp. tOV-UO, tig i 

Diagnosis: "Wide egg-shaped lorica. occasionally 

with almost circular outline; head aperture ventral. 
nearly circular, ai times bordered around lower part 
by curicular rib*; loot opening parabolic to 
elliptical; caudal margin rounded or lightly 
indented. Lorica Hat, Dorsal lorica in cross section 
.slightly arched with wide raised hump, which in 
posterior 'ird, accompanied by two converging 
pleats, carries flat keel, terminating in line poiw 
reaching caudal margin. Lateral antennae papillae 
small on end of oblique lorica fold (Fig. 6;2a)- 
Sensory pit present on distal foot segment, 

Lorica length SO^m; width 68 /*m; head apei tare 
22 f/m; lorica height 20 /inn loot aperture 23 ^tn_ 
tocs 12 pm. 

Distribution: Only known loeab'nes Mombasa. 
Kenya and Winmurra billabong, Magela Creek near 
.labiiuka, N.T 28.5°C, pH 5.4. DO 6.2 mg I ', 22 
tS cm ', alkai. 2.1 mg l '. 

I epadeliu minurut Koste 
FIG. 6:4 

L. tmmttni Kostc, 19#J. pp, 110-113, Kig. > 

D/a.L'Mosis: Lorica outline in upper pan trapezoidal 
with wings (duetlikc, tctminally concave (Fig. 6:4a, 
b)); lorica lapers .symmetrically from wings, ending 
in pointed protrusion covering foot-opening, head 
jperture (vential) smoothly indented, dorsallv wttli 
median notch; thin keel runs from notch to 
posterior lotiea margin; ventral plate flui, wiih 
robust bead-like structures (uupatierned); dorsal 
lorica covered wiih 3-6 ;*.m long rods in tows 
parallel to margins outside, and wiih keel on inner 
rows. Rod* sit on circular bases. Lateral antennae 
dorsal, located approximately level with anierioj 
margin of foot-opening, ujth pistil-like papillae; 
foot 3 . segmented; toes lighily curved vcntrally. 

Lorica length 72-80>m; width 76-80 ^m; height 
4K gnr toe length 24 flfh; head aperture width 24 
jttrili depth 12 juii; foot-opening width 16 aft, 

hi s";!, /,i'/Wl-//l/ a/tWc Hanitar $} ventral; (b) Jatcial; (c) cr^s^ecunn. 2. I Iwufunum H.irtim/. Lo veiinal, 
V0\ dorsal; kl lateral; <d) ciVKs-sceimn 3, L carnuui (tiOMfeji venlral. 4. /.. chn^uiuiht knMt- k Slu-.I; (at >cnnal; 
i\>\ UtMtit s^inciil nl I.0PI ami Iocs: (>) ncc'ipcial nmryin, Jorsal, (c) erus>--L-e[HHi. 5, /- t/wtxfiwnt (Sw\\roo>\. (,0 
venual; \ls) lareiiil; (ej eto%s-scclion- 6, /., rfccuni IJeuins; la) dorsal; (b) \enimL \ t t'lfiptua Wtillen (a) ildWftl 
OM m\W\ kj I ro'A-settiuo. M. /, Umt\hm\ tltil-.Midtiri): no vrinml, (\» laical; te) eit)»-«tfriofi> I 3 £ fc ai'iei 
llaoinuOytfOi J, aire. Kt*tv tl'Wh: I. mt,- kosic & siu.-l (Pism, 6. after Uer/ius (P.»s:»; P, ..Irei Walter! <IW>. 
Scale Jine-, ^tl it\\\. 





Distribution: Endemic, Lcichhardt and Wtamurra 
billabongs. Mageta Creek near lahiKiia, NT. 
2S.5-mi°C pH 5,4-5,5, DO 5,5-6.3 mg I », 
23-5S- & cm \ alkal. 2,7-2.9 mg I '. 

Lepadetla ocboissi Bcrzins 
FIG. fid 

Z_ fflt*r>;.w? Hemns 1*60, p. fr>, Fugs 1-3. 

Diagnosis Outline of body broadly ovale, greatest 
width slightly posterior to middle ol body; lorica 
strongly dorso-ventrally flattened, without ridges; 
anterior dorsal margin straight, ventral margin with 
wide V-shaped sinus and short fold present on each 
sade; lorica rounded posteriorly, Sinus and ibid 
slightly behind the middle of body, on each side 
of lorica, with corresponding pores on dorsal side; 
fool groove nearly afi long as wide; Foot rather short, 
terminal segment longer than first and second 
segments combined; toes relatively long, divergent, 
pointed at apex. 

Total length 105-108 &m (lorica 78-81 fim); width 
of anterior lorica 63-66 ^m, ventral sinus 11-13 >*m 
deep: foot groove 18-19x12 ^m; foot 14-16 urn; mes 
M) ,.m. 

Distribution: Fndemic, King Parrot Creek, near 
Kinglake West, Vic 

Comment: Resembles L. monodi Berlins from 
Senegal. Africa* but differs in the form of The lateral 
sinus and the pores for lateral antennae on the 

LepadeUa oralis, (Muller) 
FIG. 6:5 

Bnnhiomis ovalm Muller. I7H6, (V A45, V\^ A9;|-3. 
LefwtUila waits after Ehrsnbcrg, 1X30, p, 45, I h», 7:4 
non Metoptdkt avails Anderson & Shcpfwd. 1892, \i. ?8 
(\cc /. v'urea). 

Diagnosis: Lorica outline oval to circular; wide 
space between internal organs and lorica margins; 
ventral lorica nearly flat; lonca ar rim thinly 
capered; narrow striated area at outer margin from 
head aperture ro font-opening; outline of loot- 
opening variable; gastric glands ollcn lobed and 
long-stalked. Juvenile animals sometimes have 
caudal notch dor^ally over foot-open tng. Males 
known. Resting egg shelL covered with robust, lightly 
curved spines. 
Lorica length 90-170 ><m; width 70-140 pm; 

Tig. 6:1, (ai Lcpodella tatusinus cf. vtsenda, ventral; (b) /.. viseada Myer*. ventral; 2. / . Hndaui Kosie: (a) N.I, form,, 
dorsal, (b| ventral; (c) Mombasa form, ventral; (a) dorsal; (el cross-section. 3. /.. nehoissi Berlins: (a) dorsai; (h) 
ventral; (c) cross-section. 4, 1, tnimmtt kasie: (.i.i dorsal, ibi vent ml, (c) cu^s-scction. 5, /.. ova/is (Muller): (a) ventral; 
tfO lateral. (C) cross -section. Uj NT form, (e) Ta;». form ta. 2. 4. 5d, afier fcosie (1981); 1L\ after Mycfl ((914); 3, 
after Betzias <l%0); 5a-c. afrer Harrin* (1916). 5e. after Shirl & Kosie (1985) Scale lines 50 ufa. 

height 25-30 w m; toe length 22-36 ,un; foot opening 

to 48 urn deep, 7>2 pm wide, 

Distribution: Cosmopolitan, between maerophytcs 

in benthos of fresh and saline waters. Uncommon: 

pancontlnental. Ecotypcs described from Australian 

waters include a form from the Magcia Creek, NX 

(Fig. 6.5d) and a minute form from U. DutvcUan,. 

Tas \V\». 6:5e). R.2-24.0°C, pH 5,8-7.7, 273-3330 

uS an '. 

Literature: Ko»tc |m 1981; Shie! & Kosce |9K5; 

Kostc&Shiel 1986a,. 

Lepadelta patella (Mullen 
FIG. 7:1 

Bruchionus patella Muller, 1786, p. 341, Tig 48:15-10. 
Upttdetht patella after Dory do St Vincent. IK26 t p.S*. 

Diagnosis: Lor&fl outline ovoid to circular; dorsal 

lorica domed, without wings; ventral lonca flat, 
occasionally with somewhat overhanging margins; 
wide lorica forms more domed in cross-section than 
narrower forms (Wulfert (I960) described Hal and 
domed forms from Sphagnum): variable foot 
opening (Han« (1962) found caudal corners of 
foot-opening extended into spines, while one 
specimen described by Wulfert toe (it, had almost 
circular foot-opening); occasionally, dorsal lorica 
adjacent to foot-opening has weakly distinguishable 
folds, Male known. 

Length 120-145 /^m; lonca length 70-110 am; 
width 65-90 ;*m; length:width ratio mostly <l.2; 
toe length 20-35/an; male 110-120 pat 
Distribution: Cosmopolitan, euryopic in fresh and 
saline waters. Common: paneonrincntal. Ecotypic 
variants aie known from NT. (Fig. 7:le) arid Tas 
(Fig. 7:lg). 10.0-24.5°C, pH 3.9-H.8, DO 7.8-11.6 
mg I', 9.2-6600 f& cm\ 3.0^262 NTU. 
Literature: Kostc 1978. l'.>Kk Shiel & Koste 1985. 
Comment: Several recognized >ut>specific variants 
of L patella are recorded from Australian water*. 
The status of these tajtt may change with detailed 
systematic work; 

Lepadelta patella biloba Haucr 
FIG. 7:2 

Lepadetla patella hiloha Haucr, 1958. pp. 2?-2K, 1% 2:U. 
/ pufeila f. biloba (Haucr) b> Komc. 197S, p, (65. 

Diagnosis: l.onea outline resembles L. patella, 
however differs from f. typ. in three features: lateral 
corners of foot-opening reflexed dorsally (Fig. 7:2d); 



4f 4g 

Tig. 7;l, Lepadeila patella (Mullcr); (a) dorsal; (b) ventral; (c) lateral; (d) cross-section; (e) NT. form, ventral; (f) 
cross-section; (g) Tas. form, ventral. 2, L. patella httoba Hauer: (a) dorsal; (b) cross-section; (c) ventral; (c!) lateral; 
(e) Tas. form, ventral. 3, L patella oblonga (Ehrcnberg); (a) ventral; (b) ventral; (c) cross-section; (d-f) variations 
in fool-opening morphology. 4, L. pulelki similis (Lucks): (a) ventral; (b-d) different loriea forms, ventral; (c) cross- 
section: (f-g) foot-opening, ventral; (hi L. bttangensis Berzins, ventral, la-d. after Harnng (191(0: le, f, after Kosrc 
& Shiel (1980); Ig, 2e, after Koste & Shiel (1986): 2a-d, 4b-g, after Hauer {1958); 3a, c, after Wulfert (1939); 3b, d-f 
after Bjoiklund (1972); 4h, alter Bcrzms (1982). Scale lines 50 jim, 



ventral iorica with lateral concavities, side margins 
deflected downwards (Tig. 7:2b); two slum cuncuiar 
folds o\ caudal dorsal Inrjcn* 

Length KO-107 ^m: «vidih 59-64 fan; beighi 36 
Mm; toes 19-25 /an, 

Distribution: Europe. Rare: Tas., Vic, southwest 
W\. U,8-24.0°C,pH 3.5-7.9, IK. 3-2130 ^S cm '. 
literature: Koste et <?l t 1983; Shiel & Kostc 1985. 

U-padelia patella oblongu (Ehrenbeig) 
KJG. 7:3 

SquattUiln oblongu rhienbetg, 1834, p. 220. 
Lv/wJcfhJ obfurtvu {lihrcnbtik) &Uer Harring, (913, p. 64, 
LcfuuU-dd im/clta i\ obUmw al'lcr Walton, l%0. p, 2H5. 
Lig. 24, 

Diagnosis: Distinguished from /.. ptrfjtffa only by 
relatively low Iorica height and narrow head 
aperture. Lorica outline mostly elliptical, 

Lortea length 80-110 ^m; width 50-70 ^m; 
lengih:width ratio >1.5; toes 20-25 idft.. 
Distribution: In fresh and brackish water pools, 
often sympafric with the nominate species and L. 
patella -sitnilis. Bjorklund (1972) demonstrated 
biomelrically distinguishable populations, however 
intergiades with L. patella typ. occur. 
Only Australian record from Sheepvvash Billabong, 
near Yea, Vie, I7.iv.76 (Shiel tinpubl.). I9.9°( \ pH 
7.2 V DO 8.5 mg I ; K5 ;/S cm \ Secchi trans- 
parency 80 cm. 
literature: Kosie I97K 

Lepadelia pare/let sunifis U .licks) 
IIO. 7:4 

\fetopidia Mimtliv luck*, I9I2. p. Iiy, Fit. yt, 
LcnarMlit jfatffe tLucks) ollci Ha nor. N25, p. 1 7 
/ intfette vac. s/m/Mahei Rename. I92M, p. Fig. 1| VVK 
/.. huwincnsK lierzins. l ( *H2. p, \\ t \\t. 3 

Dtagnous. touch outline elliptical; foot-opening 
with reblfpttttd upper arch (Fig. 7:4 f, g>. mostly 
hexagonal with angled edges and base straight to 
indented. Not reliable taxonomtc character because 
or variability in foot-opening. Distinguished liom 
L fhttella fa .sir.) by Iorica lengthrwidth ratio { > L2j 
and overall smaller habit. 

Length 68-K5 /<m; width 49-60 /*m; toes 17-21 

Distribution: Heiwcen macrophytcs, detritus, in 
psauunon and rock pools; fresh, brackish and salt; 
pH to 10 (koste 1978). Single record from Aushalta 
(as L buangensis Rcrzins), hut recognizable as a 
preservation at tcl'aci of L patella sUnills. Mi Donna 
Huang, Vie 

lepadeila piilotu Berzinx 
HG 8;l 

L. piilota Reran*. I960, pp, 83-85. Figs 4-N. 

Diagnosis; Body ovoid in outline; vcniral surface 
slightly medially convex, dorsal surface very 
strongly so; dorsal median line with sharp, uneven 
longitudinal ridge; frontal margin dor.sally convex, 
with blunt projection in the middle; vcnoatly with 
rounded sinus; laterally, Iorica enlarged with three- 
pairs of thin, transparent lamellary projections; 
lorica prolonged posteriorly to rounded lobe; cross 
seclion somewhat triangular, 

Length 72-75 ^m; width 60-67 /on; ventral sinus 
depth 8 j<m; length ol loot groove 12-15 jatft toes 
12 /an. 

Distribution: bndemie. Only known locality 
Creswick Creek near CUmcs. Vicloria, Ecology 

Comment: Resembles / tripteia (khrertberg) Luil 
differs in (he form of the frontal margin of the 
lorica, the sharp dot sal ridge* and the presence of 
lateral cmicular projections Iteimcd "flappers" hi 
ihe original description although they have no 
muscular contact], 

Lepadelia quadricunnata (Steuroosr 
F-'IG. 8.2 

Mv(opidiaquadricurimauS^Vi\oo\, IK98, p. 165, tig. K. 

Diagnosis: Dorsal lorica with four short ribs ovei 
foot-opening; posterior end tapering to short 
tongue; vcniral lorica Hat; \\yot opening variable. 
Lorica length 81-92 w m: width 61-72 ^m; height 
to 34 um; toe length 20-26 /.mi. 
Distribution: Littoral, between macropuvtes. p\i 
6.2-10.2 (Koste 1978k Only record that of Berzins 
(1982) from CoghiU's Creek neat Tourcllo, Vic. /. 
<?. sexcarinofa (K lenient), recorded from the same 
locality, may be an eeotype. 
Literature: Kosie 197H r 

Lepadetlu qtunc/ueenstata (Lucks) 
PIG, 8:3 

Mctoptduuiuinquecostota Lucks, 1912. p. 126, Fig. 4? 

Diunnosis: Lorica pyriform, narrowing towards 
head; dorsal lorica with median low keel which 
begins as widely separated ridges behind collar of 
head aperture; two or three pairs of side ribs; 
distinctive cross-seel ion (Fig. #-3c)\ Joriea end 
variable, sometimes with short ridge; foot-opening 

Lorica length 92-112 /<m; uv-idth 60*70 pzq toe 
length 23-39 //in. 
Distribution. Cosmopolitan in hminal, oficn 





collected in Sphagnum pH 4.5-6.7. temperature to 
WC (Koste 1978). Single record: dam near 
Chillaftoe, Qid. 

peat swamps. Only record Ridded Creek. Sunhury. 


Literature: Koste 1978, Begins I9K2. 

Lepudella rhombotdes (GosseJ 
FIG. 8:4 

Mt'foph/iu rhombouies tiosse in Hudson & Cause. I88f> ( 

p. 108, lit*. 25:10, 

i.cpLnlfllu thotnboides alter Hnrring, 19U, p. 65. 

Diagnosis: Dorsal lorica with moderately high, 
more or less arched median keel, bounded by lateral 
grooves; anterior margin width V\ lorica length; 
dorsal sinus broadly U-shaped; ventral sinus V- 
shaped; dorsal aperture of head-opening with collar, 
which can be bordered by cuiieular thickening; foot 
opening narrow, U-shaped with parallel sides; distal 
loot segment long (cl. L etf'tptica); toe* ihort (< '/* 
body length) slightly decurved, tapering to points. 
Lorica length 1 10-120 ^m; width 55-88 ftm; tuc 
length 20-28 iiriii 

Distribution: Cosmopolitan in periphyion. pH 
tolerant. Rate: N.S.W., NIX., Tas, ( Vic. IO.S-30,7°C, 
pH 47-7 9, DO 4.6-6.0 .iug I f. 23-75(1 p& era '« 
tO 6? NTU, alkal. 2.7-2.9 mg I '. 
Continent: Two variants are known from Australia. 
described elsewhere as svp., form or var.: t*. 
thomboides haueri (Wulfert) (f-'ig.*$:5j from a slock 
dam south of Bumie, Tas. |16.5°C\ p\\ 4.7, 42 uS 
cm ) and L t/tomboides arnnata ( Dormer) (Fig. 
fc:6> from the Hrnken R, at Benalla, Vie. (27,0°C, 
pll 7 6, DO 7.0 mg l\ 52 ,*$ cm '), 
Literature: Kulikova 1970; Koste 1978; Koste ei ah 
my t Koste & Sftfe! IVKfta. 

Lepadella wiuvihurQi (Lucks) 
FIG. Kl 

Xfctoptm,, mttcnbiuxi I ucks, 1912, p. \tf % I ig. 48a, b. 
/, rotWrtbur%t after Harnng IVI3, p. 65, 

Diagnosis: Side margins of lonca cross-section 
tMend to pointed tips below level of median ventraJ 
bulge; head aperture dorso-ventrally deep, only 
ventral sinus; dorsal margin straight, without sinus: 
fool-opening variable. Resembles /.. davfyfiseta, 
Can be confused with Colurclla spp. in lateral view, 
Lorica length 75 /art; width 50 /*m; toes 25-28 /mi. 
Distribution: Isolated occurrences in Sphagnum and 

Lepadelia (ana Koste & $hki 
HG. 9:2 

Upadcitu tumi Kosie & ShicI, I9&7, p. 102, Hg. 19. 

Diagnosis: Lonca btoadly ovoid, ventral lorica Hal, 
dorsal medially convex in median cross-section.: 
dorsal margin of head-opening almost straight, 
ventral margin with weak V-shaped sinus; fold 
behind middle o( body on ventral surface, with 
pointed, slightly curved spine on either side, directed 
apieally; foot-opening broad, toes relatively long, 
straight, sharply pointed, 

Lorica length 70 ptBJ width (medially) 61 um\ 
anterior width 29 /ur*; loot-opening 18x18 /tm; toes 
29 /mi; lateral spine length 18 «m. 
Distribution: Endemic, only known locality L, 
Pedder. Tasmania. I6.5*C. pH 5.3, 46.2 /tS car'. 
Comment: Resembles L nefroissi. a Victorian 
endemic (Fig. 6:,Vi, however the latter has ventral 
folds (no spines), and differs in foot and tew 

Lepade/ta triba Myers. 
FIG. 9f3 

Lvpadetia tnha Myers. 1934. pp. 4-5. Fig* 10-12. 

Diagnosis: Lorica elongated; erosvseciion shallow 1 . 
evenly arched dorsally; dorsal margin ol head 
aperture almost straight, venlrallv* broadly V- 
shaped; no stippled collar; ventral lorica with 
distinctive t'uch/anis-Wke slightly elevated mid- 
section (Fig. 9:3c). lateral margins run to head 
aperture or diverge to side apices; foot opening 
daring; distal foot segment approximately ! /: toe- 
length; toes long, tapering to slender drawn-out tips. 
I oriea length 71-80 (im; width 42-15 /j,m; ventral 
sinus depth 12 /uu; foot opening depth 20 /m»; foot 
length 2b *un; distal segment 14 /an; toes 30-33 /un 
Distribution: North America. Single record from 
QUI (Berzins 1982) needs verification. 

ijepadtfita snptera {t-hnMihcrtO 
FIG. 9:4 

Metopidtct (ripteru hhrenberg, 1830, pp. 7 
LepoUeflv tnptetv Ehrenberg, 1832, p; 72, 

. 74-83, 

fig. K I, UpadeUa ptilota Bemns: ta) dpftflfc (b) ventral; (c) crow section. 2, /.. quadricurinuta (Stenroos>: (a) doraaj' 
(h| vrniral; (e) cross-.scc don; Ul> dorsal; (e) ventral; (f) doisal; (t;) cnus-scction. 3, L. ytmupweostuta (I .ueks): (;0 
vcniral; fb) lateral; («) eross-scclion: (d) ilorsaL 4, L. rhamboUies tOo.sse); (a) vcmral; (b) lateial; (c) dursal; (d) cro-ss 
»ecCuom 3, L. rhomboiUes iuiiMt (Wtiltcn): (a) NT. form, dorsal; (b) l.uropoan form, vcmral; (c) cross-sccfion. f», 
f thumhaittes < atthaiu (Doitneri: (at central; (b) Jaieral; (c) wmral; till L*ft4s section spec. U! (ef ctoss-seciion spec 
•:: ti) l-.Mcn stHhice iiMirp|niIoi;.v. I. alicr Bcivhv, (I960). 2a-c, I; 5h. o, ahci B.ti.v^ (tsiS 1 *): 2d, g, alier t\-jlcr (lOh^j, 
t-\ .um Kjo.kluitd (t972); }, 4, after Harniv. (1916); 5. idler ko>le i)98l»; h, nU?\ Oonner (PM3). Sc.le imc* 3U^«l, 





Diagnosis; Lorica outline variable; circular rn 
rhombic according to Koste (197S). Dorsal lorica 
vaulted, with high, somewhat protrudine median 
keel; head apetfure with shallow U-shaped sinus 
wnirally, usually slight median notch dotsally; 
'erica surface may be finely striped (Fig. 9:41} wirh 
small rounded protrusions, most, however, are 

Lorica lengrJi 58-65 urn; width 50-57 /itm toes 
13-22 M m. 

Distribution: Cosmopolitan rn fresh and brackish 
waters, and on muddy littorals. Rare. N.S.W., NT., 
Qtd, 7av. Vic I2.4~20.0°C, pH 4>-7 7, J7-333U 
fiS cm '. 
Literature: Shiel & KftftC »9^s>; Koste 1981; Rerzins 

Lepodetlu tvleri KoMe A Shiel 
FIG. 9,5 

Lcpudrtta tyleri Koste & Shiel. iyH7a, p, 37, Figs 3, A. 

Dynasts: Rigid lorica Of trape/otdai outline; cross- 
sechnn nearly triaugulat; dorsal plate with median 
keel; dorsal and ventral plates finely granulated; 
head aperture lightly convex dorsally, deep V-shapcd 
Sinus ventrally, with lateral lamella; caudal lorica 
with slight constriction above blunt lateral corners* 
fool-opening broadly flared; Three foot *cgmeim, 
distal with caudal antenna; toes very thin, 
elongated, pointed. 

fetal length 123-125 ,/m; lorica length 78-SDurn; 
greatest width 58 /<m; height 36 /im; head aperture 
width 21 ^m, depth 22 /Am; foot opening IS\ISimi; 
distal foot segment 12 *im; Toe* 32 /j,m. 
Distribution: Endemic; only known locality a stock 
dam south of Swansea, east coast of Tasmania. 
13_<y>C.\ pH 6.0, 390 ,/S cm : , 82 NTU 

Lepudetla vttrea (Shcphard) 
VIG. ${ 

Meiupiditt ovaiis Shephard in Anderson A. ShrjiliaiU. IH*J2. 

p, 7S, FJjJ. 6, 6n ihm \f. kyifis <\tuller, I786j 
M, vittvQ Shephard, (V1K p 55 
Lcpudetla vi/rca after Harrinj!. l*>J3 f p. f£. 

Diagnosis; Ovoid, transparent lorica, tapering to 
each end from widest part posterior to midline: 
head aperture slightly convex dorsally. without 

sinus, ventrally with deep, broadly elliptical sinus; 
dorsal margin of toot opening convex between 
lateral points of slightly outcurved caudal lonca 
projections; ventral foot opening deep '^-ellipse: 
disLal foot segment almost twice lenglh of proximal 
segments, more fhan half temrth of ioes, toes short, 
tapering to pointed tips. 

Lenglh 125 pQ\ width if* ,<m. [No oihet 
measurements given in ihe original description, 
however from Fig. 6 in Anderson & Sbcphard 
(18921, on ihe basis of their length/width 
measurements* the following :ire estimated: head 
aperture 23 /an wide v 20 /<m deep; foot opening 
30 finx wide k 22 fim deep; distal toe joint 12 ^m; 
toes 17 ^m.j. 

Distribution: Subsequent to the original find 
I Brighton, Victoria), L. vitrea was recorded from 
South Westbnd. N.Z. (Russell 1954), Dimensions 
were slightly different (114 /<m long, loci 20 /mi). 
with "a series of dots round the lateral edge of the 
dorsal plate". /.. vttreu is possibly more widespread. 
Comment: /.. vitrea was synonymised with L 
htusinus by later reviewers (e.g. Koste 1978), 
however can be distinguished from the latter b\ 
more elongated lorica <length:vwdrh ratio 2.2 vs 
<1.5 in L. iutusmus), absence of cowl-like head 
aperture, elongated distal loo.* segment and 
markedly shorter toes. 

l.cpadelfa wdliamsi Koste & Shiel 
HO. 9:7 

l.tnadcllu wiliiumsi KnstcA Shiel, 1980, pp. 138-139, ft& 

Otaxnosts: Smoothly oval, highly vaulted lorica, 
with vent rally-directed head-opening; dorsal lorica 
tapers caudally to variable-length laterally curved 
pomt; foot -opening narrow; ending at base of dorsal 
lorica projection; three loot segments of similar 
length; long, doiso-laterally curved sharp toes. 

Lorica lenglh 112-116 /jh; widih 60-64 f iw; 
height to 56 flirt; head aperture i3-36s28-30 ^mr 
caudal lorica projections 19-2* /mi; foot opening 
17-20x19-20 /on, distal foot segment 10 pu\\ toe 
length 36-40 ;uti. 

Distribution: Kuala Lumpur, Malaysia, and Jabi 
Uika, NX. only known localities. 25.5°C, pH 6.2, 
DO 2.9 tng I \ 62 p Scm ', 5 NIL. 

Hg M:i ( fepade/h rottenhurpj (I ucks)r <si) venmdj tb) Intend; &_') ventral; (c\) cross-section 2, J . tuna Koste & Shiel, 
ventral. 3, L trihu Myers (a) dorsal; (b) ventral; (c) cross-section. 4, / , trtntx-w (RtrenbeoO: (a) dorsal; (b) ventral; 
[c) ernv-seaion, <d) uoss-hecium; te» lateral; (I'-h) Jiflercnr (orlcft hit rwto ^niial; (i) Tus. form, vciural; tj) 2nd lu\ 
form, ifiitinl. '\ t lyk'N Kasre & Shiel: (a), (h); (c) laieral «U cross-section. 6. L. vitrea (Shephard): 
(a) dnrstft; (h) vennal. 7, /.. uiltiatmi Koste & .Shn;l; tu, b) (wo morphv. veitiial; (c) iuieral; (d) crowsecrion. lie, b 
;oicr Ollteg 0*39); lc, d, (ilier Hanos (1059), 2, 4t. j. nfler Kosru & Shiel t\WVr. J, afier M>ers (193-1); 4a )i ( (jftcj 
1 tiRIVina (I916); 5, atici KOttte^i Stiirl (|Vtil74k &*- oria.; 6b, aher Shuphrtid itstll); 7, after Ko»W AiShM il'^D) Settle 
lines <() /mi. 


key to species of (In* genus Lt-pudettu 
(Heterolepiuielfo I 

I . I mica ovii I 2 

Innert rhonthoicial .....? 

2{\\. Head apciturc wiih V-shapal dorsal attO vetilial 

sinuses. L til), hciurtnhuiylQ Fadcev 

Head upetiure wad nuiv a veutial sinus, dorsnl 
iturtmi straight L. (H.) upsivoru Myers 

3U). Loricu Willi poskotlakTuI pointed spuis; loot {-move 

with luleral pointed piujeciions 

, , / . (II, } chrvnhvrv.i (I'eny) 

I aleral wings (vf lnnen wtU pusiei mm margins pf fool 
groove voniukil /.. til) heturosivki Mm ray 

Lepadeila (Heterolepade/ktj opskora Myers 
1IG. io:l 

/ eparfetla op.sicum Myers, 193-1, pp. 5, 7, Figs I6-1S. 

DictMtosis; Lorica o\al; cross section shallow, evenly 
arched dorsally; dorsal anterior margin nearly 
straight, venlral margin with deep V-.sha.ped sinus; 
slipplcd collar present; foot groove wide, flared 
posteriorly; distal loot joint 2\ length of combined 
first and second joints; \'oo[ twisted, left toe lies 
beneath ric-hl; iocs asymmetric, right 2\ length of 

I orica length St) (iM; width 65 /mi; ventral sinus 
depth IS />m; anterior points 28 ^m: foot gioove 
2l ^m; foot 30 /*m; distal segment 20 /<in; right loe 
mpto; left u v m. 

Distribution: North America, Two records from 
Australia: L. Boon, central Victoria, and Winiruirra 
lihllaboug, Magela Creek, NX 23.5°C, pH 5.4-7. 9,. 
DO 6 0-6.2 mg I \ 22-150 ^S cm , 67 Nil, 
alkal. 2.7 mp I '. 
/ itemture: Koste & Shiel 1980; Koste 19X1. 

L OH ehrerthcrgi (l'crlv) 
TIG. 10:2 

Nuttiwnht t'hrvnhemi Perty, 1850, p. 20. 
Metaptdta ehtvnherp otter Jennings, 1S94, p, 2fS. 
f fj>Ltfl?l}a riircnhergt alter Harrinu, L9J?, p. ftfl 

Diagnosis: Lorica rhombic, with posterolateral 
margins produced lo triangular, dorsally ridged 
spurs reflexed upwards in cross-section (Fie. 10:2c); 
smaller pointed spurs on each side of fool groove; 
stippled collar on vemnil plate, with less obvious 
dotsal stippling; fool groove *A length of loiica, 
rounded anteriorly, flaring slighlly posteriorly; fool 
stout, ,j 3 length of lorica; distal foot joint longest; 
Iocs long, asymmetric, taper evenly. 

lorica length 70-95 ^m; width 70-90 /*rn; loe 
tengtii 27-32 ^m and 19-27 ^m.. 
Distribution: Cosmopolilau in ]>enphyion v (•'tricn 


lartu and moss. Rare: in billabongs and vegetated 

lake margins in N.S.W,, NT., Qld, Vic 24.0-29.2°C 1 

pH 6.3-7.5, DO 5.1-8.0 mg I ; , 42-85 ,iS cm ' j 6 


Literature: Shiel & Koste 1979; Koste & Shiel 1980; 

Green 1981; Koste 1981. 

L. (H.) heterodactvla Fadeev 
FIG, 10:3 

L (H.) hcterodauyia Fadeev. 1925, p. 73, Pi. 1, I \$. H. 

Diagnosis: Ulrica oval; dorsal and ventral margins 
of head aperture wilh deep, almost triangular 
sinuses; last foot segment longer than preceding 

1 orica length 106-U3 /tin; width 70-75 /tm; distal 
foot segment 22-23 /Am; toes 23-30 and 20-25 jtm. 
Distribution: Furope, S. America (Amazonia I 
Single record, 1.. fioort, central Vic. 20.0 *C, pH 
7.4, DO K.O mg I '. 

Litcraiure: Kutikov;i 1970; Koste 197k; Koste & Shiel 

/, HI) hetenwylu (Murray) 
NO. 10:4 

Meiofudia heteroslykt Murray. I9I.V p. 459, W. I'J, fig. 


Diagnosis: Lorica rhombic; lateral wings rounded 

lather than pointed (ef. /_, ehrenhergi), with tips 

more or less deflected dorsally; ventral lorica wider 

than dorsal m region of head aperture; stippled 

collar present. 

Distribution: Probably cosmopolitan in littoral 

/ones, stagnant wuters. Rare: N.S.W., Tas., Vic. 

10.0-23. 5°C, pH 5.8-7.9, DO 6.0-9.8 mg 1 , 

80-750 pS cm \ 2-67 NTU. 

/ '. iterative; Koste 1978; Shiel & Koste 1979; Koste 

& Shiel 19K0. 

Not recorded from Australia: 

L. (H.) cyrtopus (Hairing). Eastern Europe, N. and 
Centra! America. 

A single free-living species of ihe subgenus 
Lepadelia (Xenolepadrlla) is recorded from 
Australia. In view of the preferred habitat of most 
of the other known species of the genus, i.e. the 
branchial chambers of a range of crustaceans, it is 
tikcly that other species occur here but have been 

i.epadeltu (Xenolepadellu) mondactvla Bei/ins 
] Ki. 10:5 

1 4'PiJiidia tiwnor/actyla Bemns, CJfiO, pp. 5, A, Figs R-ll , 




Fig. 10:1, Heterolepadella upsicora Myers: (a) dorsal; (b) venlrat; (c) eross-seciion; 2, H. ehrenberyi <Perty)- (a) dorsal 
b) ventra 1; ft) cross-section; <d) lateral 3, hi heteroclnayfa Fadeev: (a) \entral: (b) toe detail. 4. //. hett>msn>/u Muirav 
(a) ventra ; (b) dorsal; fcj lateral; (d, e) different cross-seciions: 5, Xenole^ideUa nwnodacivta Herrim:" (a) dorsal 
S2JS5 I, ?'S iCy»MWee|»n- I. 2, after Harring H9J6|; 3, after Kutlkova (1970); 4, alter Harrine <19lfi): \ nller Her/his 
(l%0) Scale lines 50 pm, 

Diagnosis: Ixniea outline elliptical; dorsal lorica 
arched, punctate or lined; ventral plate with light 
bulge medially, otherwise unstructured; head 
aperture with V-shaped dorsal and ventral sinuses. 
Total length 170 /mi; lorica length 127 /mi; width 
77 jim; foot-opeiung 31x10 /mi; toe 57 /tm. 
Distribution: Madagascar, variants known from 
Brazil. Single record, billabong, Jabiluka, NT. 
25.5°C, pH 6.2, DO 2.9 mg 1 ', 62 fiS cm f . 
Literature: Koste 1978; Koste & Shiel 1980. 

Not recorded from Australia: 

L (X.) astackola Harring, L. (X) borealis 
Harring, L. (X.) branchicola Haucr, L, (X.) haueri 

Rodewald, /". fA^fa/a Wisniewski, L. (X.) parasitica 
Hauer, L (X.) pygmaea (Gosse). See Koste 
1978:198-200 for details). 


Collectors acknowledged in our earlier papers 
also contributed further material to this work. Their 
assistance is appreciated. The Deutshcen For- 
schungsgemeinschaft, Bonn-Bad Godesburg., 
provided microscope and photographic facilities to 


VV KUSU- a I. 5HlH 


AMI ,ik»»M. I H |I9<M I'lankioti Rotatoria ihflW Nurtlt 

Catulrrw, J frWtytf KUUhrll S.7 Sot. 54, KK-110 
\itnAi"-, n, <I957| UuDiNUKli-OkoloiitM h Stildten an 

Koiairn ien snlt/ftjliuni Gewu.s>ei MiweUkut*>ehlArtrts. 

Hi vs. Z. Murttn-! urht>t~t'nt\\ Uutfo W it U'tiherg outfit. 

twuiruMs. 6, 1(7-58. 
ANor^soN, M l( A. Mhi'hakd. i. (18921 Noiis on 

Victorian routc'v ptoe. A'. $}>r, IV/. 4, h9-80. 
Hakio-., I . (1955} Viiil'Ci ttftlju / rywr/Wto U W& K nmt It 

urcmvni, Hff&rf \annln, Wt £*tfrf /wfrwtf- 124. 

(1959) Virtue) — Romtorij. /wwa C A & J5, 

BEK7IN--:, K (1960) PflCDti Kotaiorieti EtUS MatUtitrtikar: 

Stcfti, Hh&h SCr, MiM S*f A. 14, 1-6. 
(1982) Contribution Hi the knowledge pi KoTatona 

Of Auvitalm (UnivcrsiTV of I und, Utnd) 
HHWkM<M», Ii. 0, (197,1) The miitcr tauna of ilic rock 

p<>i>i <• imiu'Tvarminnc Arehipelu&it. southern finland. 

Ada /»<>* t-ftn). 135, 1-30. 
Hum ix Si VirfONT, J.-&. (Q, M. M (1*24) £fcpj>. 

flS2f»i l&sw d\we cla^iiKJiion tlc» animate 

niieroscopk|ue;' (P£irf\> 

C\riin, \y (-1039) (Jhcr koiuionrn eimyei Seen bci 

Aitebody. vre>/<y. f imnoj. fust. 1 u/nh fjtiiv 2, ?-$8> 

< \ -a>n, B. (193^) Hhcr die iinieen fur Sehwcdeti 

neuil Kntdiifrien, /|r& Zw/. 26A, l-14 ( 
OtlftUAj A- P. I <I8JR> S'ouvelles observation?. 

it'ifOKopiqucs. Atmunuch de CurhfradM* pu-jyg. 
PKOAVi t* Via H9()tfi Adatok Nemet-Kck'S-Alitlo 

cvle»A«?T mikiVviminajMnak i-anereiehei. {\t3sodtk 

koitemcnv.) ihid 26, 1-42. 
Duvmk, I. (19431 Zur Rotalorienlauua Smlrn-iiii'cus. 

/W d«2 143, 172-179. 

(l*>5lj Rotaioneii tier Hnmusboden. HI. Zftti 

u/ir/>. ,4/>/. S'VAf. 79, 614-638. 
Dubois., F (IK4t) 'Hisiotre naturcllc dc*. iuophyies 

inftp-oires eomprcnaw la physiblo£itJ ci Ui manure Ac 

les cutttifi rt I'muIc du micrxweopt 1 . (Pan>.) 
Timi SJJiiHO, C C. (IX3(t) C)r£;-iiirii»ijon, SvMt'aKilik und 

tjci^)iiphi,Si.'hf.\ Vcrhffl.lJBwJ tier inl'uMoiisitiifiehui, Ahh. 

Ahtid, Hns. fieri- i»V.r I82S and I«30), JJp. 1-108. 
(II3J) Hemprivh el LJirenlKig ,Syniht"»bephyiicue. 

Anminha evt:nc*hraia, e\-.lusi^ hisectii, pCK'CrtMiir l)r 

i. . C Hirnihete, (Horolini) ( lt\i unpagcxlj 

(J832) /i"' 1'rkeniiinis der Organisation in dcr 

Rii'huinK ^it- ,k . kk'in.sten Raumes 2. l-nrwicketuny, 

tebcifvlMLiei' nnd Mniciur der Vt;uteiiilHen: und 

Raderllncce. 'tt.lci ^rtueniininen Inl'iuorien. nebit einet 

^hysiologisehenCtia'iaeteMMik heid«r Klrtwen und U2 

Arret) dtfffict&en. .1/Vi. /1A^/. I''iiy AM Cot »X^I). pp. 

UK.**-)) Deiisa;.* ? '" r-ik'.-ntiTitm wrosser Oruaaisalion 

to dev Kii.lnanude''. kleiiiMeti Ruunto 1 IhuHkn I83J), 

R p, 145-^36 
f RikM n, B. (i. U%8l Mitftite mritor^ found in Norway, 

\'.tth JesLfipnons of imt new and oti\* lit^tc known 

\peeies. Sitisitttt* %% \U 
t \'.s-., I. (i95U Tl'L totiici letaid oj' Vieioria. /V(;r ( 

If/^mw Stftf. IV(7., -1—7. 
luitcv. N |IM)?J Vlarerialen /ui RoiafoiL-u-latmn tics 

SSSR, Kwue diaj'.ttosi.'i» neuer Kounorien der rulfiibCh^n 

l.iuniL ftussk. Gidruhfah Sh. 4, 72-74. 
KRr.r : MH\, I. H. O. \S. (IK58) Bvlti&V '"« Konntni). 

miKtoskoinstiti Oivan^mcn, Ahh. Sertckvirfi tyt <iVh. 

2, 211 242. 
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NW/uni lijitsihi. Ml isv 2ia. 

en /vsim r, i.s. (i»9.n a i'U) cd sonwey Phc Roi»ter*of 

IfdOtnU- &•! f*Y«. W. /JtfWm Sot: MS. ». ?5-W 
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J h. Micmtt. Soc^, 1-7. 
t4:fr\. I. (1981) AisatfjdtqmS of roiilcis «n Au^iratKirt 

crater lakci / ZbOh |L">iui.| IW, 4^;*-jh6. 
Hahkinc, H. K. <I9J3'i S\iu>pm> o1 the Koiaiona. Bull 

Am Mux: «». l-22f»- 
_ (1916) A revisuiti t»f ifu mi.itorian genera 

Lepadcth and Lophochurib Willi dcscnpitons o( five 

tu?w spceics. ftrwc (.(5. .V«/ \fuy 51, ^27-^f»H. 
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(1925* Rotaiorien aus der Sat/L-ewasbern von 

Oldo>ioe (HolM) $4tir, Owgr. fhf\) Nat. Uhi. May 

ItiheCk 30. 152-195. 
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(Oouland) und etniiieti andern ^crrw>jdi-chcn Hinnen 

gewasssrn. Ark, Toot 6. 1-251 
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(I ongman, London.^ 
Hi T(HiNJsoN t G, E, (1967) A Treatise on Limnology. VV»1. 

2. (Wiley, N.Y.) 
Jt'-JMNas, H. S. (IKy4) A \U ol l)ic Rotaioria of rjbc 

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Kumi , W. (iy62) iJbcr uic Radeiheitmnm d«*s Onfnscfr.* 

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'Waturw. \t:r. Osauhruck 3tl. "7>- 1 37, 
(I972t Kotatorien aus Oewaasetn Aina/oniftts 

Amuzonunui 3. 25S-505 
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(IVSf>at New Rotifeia (AsetKMminrh^i from 

'lasmania. Ihirj 110, 93 109. 

(I98hb) Rufitt:ifi hv>m Ausiralian inUud waicri. 

1. Bdelloida (Rolifera; Digononrrt). tirWiU ./. Mar 
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_ H9S7a) TaMnanirin Koiifera; afilniliei wftfl QW 

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& Kn, L, W ||9«8) N^m ro|ttci> (KoTtUv.-t) 

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Nauk. CCCP, Leningrad. ) IRu^sian.j 
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Wulfert, K. (1939) Bcitrage zur Kenmnis der Rader- 
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(1950) Radeniere. Naturschutzgcbiet auf dern 

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by A. D. Short*, R. C. Buckley**, & D. G. Fotheringham'J' 


Field investigations along 2150 kin of the South Australian coast including the Great Australian 
Bight, Eyre Peninsula and Kangaroo Island revealed numerous sites with low wave energy beach 
ridge and occasional chenier plains. Examination of 14 mainland sites and six sites on Kangaroo 
Island indicate that all ridges are wash deposited during episodic wave events, probably over a 
period of decades to centuries. The ridges overly intertidal shelly sandflats. The oldest innermost 
lidal flats date at 6.3 ka and 5.9 ka BP, but the oldest ridge dated at 4 ka BP suggests a period of 
tidal flat formation and progradation is required before a source of sediment and plat form for ridge 
formation is available. All ridges and ridge sequences dated between 4 and 2 ka BP with no ridges 
material younger than 1.8 ka BP. While this confirms a delayed start for ridge development it does 
not necessarily imply no present ridge building as reworked, older shells may be incorporated in the 
modern ridges. The age and elevation of the sites lend support to a published sea-level curve for 
northern Spencer Gulf. The limited evidence available suggests that sea-level reached its present 
level in the Bight by 6 ka BP with no conclusive evidence for changes since. 
KEY WORDS: Beach ridges, sea- level, Eyre Peninsula, Kangaroo Island. 

thtriwi-fiont ihi' Hoyaf Society of S. Ami, (1989), IM(0 145-ttf. 


by A. D. Shori*. R. C. Bt( ki i \*\ & D, G. Fothi kim.ham I 


Mioio. A. J)-, Hi * mi v, H. (\ & Foiur«won*vi, D, <> (IsW) Frelimmaiv imcMipalious of beach ridpe 
nrogtadalion o\\ Eyre Peninsula and Kangaroo Island, Iran*. R. Soc. S. AuU. 1 13(3). 145-161, 30 November, 


I'K-U imcshualKUVs alonp 2(50 km ol (hi. 1 South Australian Coast in-eJudtUy tlHJ ( nc:tt AiislraUan UieM. 
I£wv I'cniusiila jfifjil Kangaroo Island revealed utuneruus sites with low wave enog) beach fidtfc and occasional 
ehenioi plains, LvAininaHon ol 14 mainland sites and six sites on Kangaroo Island indicate dim nil rtdfjes 
an; tttaMi deposited during episodic wave events probablv over a period of decades ro centimes i lie nde-cv 
over's iuicuidal shclK sandl'Kus I ru oldest imuYORVxi JidiO Mais tluK* at M itij and 5$ la UP; bill (lie 
nldeM udye dated ui -1 ka HI' Slfltftw a peiiod of lidal llal fornruuon and prouiadatinn is ieo,inied l?fifwti 
a somce of u'dirucnt and platform for ridge formation is available All ridges and rT^gc -equences dated 
between 4 ami 2 \m liP with nn ridues imiienal voiiiikci dtan 1.8 ka BK Whdc this i*dt»/ijTtW( n delayed 
stfut lot 'idue development it i\o:\ not neccssirilv impiv no present ricttie biiildinu as reworked, older Mielts 
may he incorporated in the modern ricivc-. I ho age rmd elevation <>l tin/ sites lend support to a published 
mm- level iuive lor mtiihcin SpfMl J £t Oulr", The limited evidence available surest s that sea-level reached 
its piesciit level (it "ho Hiyln hi" & Ka BP \Wrrt nc> eonchishc evidence :)or changes since. 
frCrv Words Beach ridges, .sea-level, Pyre Peninsula, Kangaroo Island. 


I lie South Australian coast, west of Whyalla and 
including Kangaroo Island and the 7S islands of the 
Eyre Peninsula, represents 60- of the state's 
coast lute. I lie coasi contains a variety ol I loloccne 
shoreline types ranjiitm from tide dominated 
suudflals lo, with increasing Uigyc enemy. ehcnieis r 
beach ridges and regressive (lorecliirie ridges) and 
rrans^ressive banicr-dune systems (Short IWSa). 
I he latter include some ol Au.slraliaV most extensive 
coastal dune systems, including more llian 500 km 
of clilitop dunes (Shod &t$bh 

Whilsl the .^700 km .S. Ausl. coast possesses a 
v\ ide [ttfl&V of I lolocene depositional 
systems, the iiautte and evolution of these systems 
received only cursoty attention uniil (his decade. 
Regional studies and detailed investigation have 
onlv just begun lo address Hie status ol the coasi, 
In the euir reuions HoIocciil* sedimemaiion and 
Pleistocene and Holoeene sea-level histoi v 
associated with beach rtdce plains have been 
investigated by Rurne <I9N2>, Belperio ft <//. (p)8}J, 
Haiis & Gostfll EW54) and Helperto tv id iV)HH). 
IIailscv(ioslui (19X4) and colleagues also presented 
n series of papers on the evolution of northern 
Spencer Oult and its imphL-aiions I'or modern 
scdimenlation. In Sr Vincent Gulf, Bowman Sc 
Hancv (I^S^i conducted extensive coring and 

■ < oaual Studies t.'nit, Dvpl W Cuojuaphv. L'iuver>ilv 

ot SsdfiL-v, Svduev.. N.S.VV. 2fl(Jn. 
•• AMOtii. I A3 Aiillci Si, C)'( onnei. ATI 26Df>, 
I ( oa>i Manamaiieni Bran- It. IXL.I 1 .. IM). bo.\ (^, 

Adrla.df, s, \wsl 5001. 

datiniiot' the I e Fevre Peiiinsula lo reconstruct tlu 
cvolutmn ol this iinptutaiH heach-itdye coniplcx. 

On the open coas| Ihree rectml regional studies. 
of the South Fast. Kanrjaroo Island nm\ Eyre 
Peninsula tShon & Hesp 1 9H4; Shot! & 
h'lhciiiiiiham l l ^6; Slu.irt ci ttl. I^fe) (Movidc the 
Hi si eoinpiehensive asscssmeni ol tlie miupho 
dynamics and Kolocene evoluiiott (if" these sections 
^)f coastline. The aim <jf this paper is to assess tlie 
evolution of the low wave energy beach ndge and 
chenicr sites on Kant'atoo Island and Fyrc 
;md their implications for mid to late Hotocene sea- 
level in the reeion. Heach ridges ate heie defined 
as shore linear, low sandy-shelly swash deposits, 
separated by widet iittci and supiu-tidal sand Hals. 
Ihey are a pioduct of periodic wave action and 
should not be confused with aeoltan fotedune 
ridges, which have also been called beach ridges (see- 
Hesp I*)X4). Cheniers have similar ridges but ace 
utldei lain and separated hv finer sediments .such as 
muds, slits or I'mc muddv sands. 

Heaeh ridfcs occupy only 4" (66 \im) of Fvrc 
Peninsula and (,(i"< ) 30 Kan of kangaroo Island. Tins 
is run .surpiisiug j;iven the generally hii'h wave 
energy together vviib hmh percentage 0) WKkv coast 
(6f> ai>d &% aspectivcly), On Fyie l*et\insula beach 
ridges ate common in the cenlial gull and occur 
in several ptoLeeted bays on the Bii^ht coast, while 
on Kangaroo Island ;hev are icsiricted to the 
protected ninth eastern coves (see Short et ul. P'86 
and Short & lotheringhani l'>8d lot Uieaiionol 'all 
silcs). I hesc sites are, however, \vrx important for 
the mlerprelalton oY bt^h I lolocene shoieline 
evolution and sea-le\el historv. fiaiuculatly jjtvvti 


A. D. SHORT. R. C. DLMlKLliY & P G. MriHhklNCiKAU 

the noise or absence (due. (n erosion) of the more 
dominant higher energy beach and dune deposits. 
Httih Hume (J0R2) and Bclpcrio er trf, (IJNUTf KV8) 
use such deposits to interpret Holoeeue sea-level 
history, patiicularly in northern Spencer Gull', while 
Burne tittOQ and Uowman & Haivcy UVNb) 
identified episodes of riJgc building in northern 
Spencer and St Vincent gulfe respectively, This 
paper provides a preliminary assessment ol ihe 
initiation and timing of tidge building at 20 sites 
together with some implications for mid to late 
Holocene sea-level in these regions, 

Sfuil> Area 

All head; rid^e-chciiicr sites were identified by 
mapping the entire coastal gcomorpholoe-v using 
l:U) T 0(><7 colour aerial phoiography. Twenty 
representative sites were selected tor field 
Investigations *'g- W. They represent about 50°/o 
ol both regions beach ridge systems. Table 1 list:* 
environmental parameters ol" each site. 

Field investigations involved surveying ol cross 
riilgfi transects using a theodolite, and both surface 
and shallow { < ! m) subsurface sampling, ihe laiiei 
using both shovel and hand auger. All samples were 
■collected within 10 cm slratigraphic units usually 
from in $}lU tidal flat deposits. All shells were 
collected wuhin each layer For dating only the most 
intact valve or bivalve shells were collected assuming, 
that the more intact and fresh the appearance, the 
less distance they have been lianspoitcd and 
therefore the younger thev might be, thereby 
providing a moic accurate minimum age of the 
deposit, l.nless otherwise indicated, dated shell* 
were obtained from th<c base of the auger-shovel 
huk I he location of each sample is illustrated m 
the accompanying figures. 

All sues are characterised by low breaker wave 
conditions, with essentially zero modal ocean wave 
effects Wave energy is delivered both by greatly 
attenuated { < I m) extreme ocean waves and, more 
frajuentty, by local wind waves t<<l m). Tides 
Hingp Fn *rn micTo ( - I nw on the open coasL 10 2.5 
ui on the mid Spencer Gulf (Tig. IF Orieniation 
lo the dominant west to south-west winds ,ind sea 
breezes* i-, important onlv m t elation to wind wave 
veneration, since the heach ridges sires have Imte 
or no aeithan eappu'eu. 


Spencer Citili 

Fhe live Spencer Chill sites are listed it) Tabic I 
Und illustrated in I ig. 1. I idal range incicascs from 
I in at lulka Well to 2.5 m at Olensea. OTerisen and 
lulka Well receive pel iodic low wind waves, while 

Pojiu Co mein. The Knob and particularly I ipson 
Cove receive tnore regular somberly wind waves. 

Vfenseo ' Bern h Ridges 

The eight swash deposited beach ridges (site I, 
fig. 1; Fig. 2) near Glensca Homestead are pan of 
an extensive beach ridge plain which runs 110 km 
from just south ol Whvuila to lilekV Bay. I he are fronted by tviensive imcriidal s;uidflai\ 
totalling W0 km-. Many of the ridges, such as 
those at (tlensea, iutcrdigitaie Willi pailially 
submerged Quaternary longitudinal dunes, with the 
dune.s forming low "headlands' and ridges occupying 
the intervening 'embaymems\ Dating of five o\' ihe 
ridges (Fig. 3a and Table 2) suggests ridge 
development commenced after 2.6 ka HP. The 
mixture of a$es in ridges 6, 7 and 8 suggests that 
Ihe cutler lidges may be contaminated by older 
shells. If so, their -age may be younger than 
indicated. Outing this iirnc their carbonate content 
increased from 2 to 3CM>O a /o indicating dependence 
on biogenic production for sediment supply. The 
surface slope of the tidges and particularly the 
swales (Fig. 3a) could be attributed either to a fall 
in sea-level of 1.0 in during this period, or a 
reduction in incident wave energy because nf 
sandflat development, hxamination of the 
subsurface rnorphONtraography will be requited to 
assess the causes. 

point Germein Beach — Fanx/une Rutges 

Point Germein (site 2, Fig. 1) is part of the 13 
km long VVindmdl beach-barrier that has progradcsJ 
northward, thus partially blocking 1 laukliu 
I larbour, and subsei|uemly seaward. The age ot ihe 
innermost beach ridge ai i he tiauseet locaiion only 
indicates the arrival of the northward prograding 
banier there at 3220 2 80 yr BP (Fie, to; Tabic 
2). Subsequently, the banier has viogi acted 250 111 
eastward into the Gulf wilh increasing acoliau 
capping producing a 5 m high foredune at (be 
preseni shoreline. 1 he elevations ol the three inner 
beach ridges are all wit bin the hunt ol present day 
-.wash action and also surest rapid shoreline 
progradaiion compared to the higher outer ridges 
(4-8. Fig. .lb). 

The Ktwh — Sljingb Pktgex 

The II exposed shingle ridges at I he Knob (site 
3, Fig. 1; Fig. 3e) record siurm wave deposition of 
coarse Tertiary conglomerate material. Whilst no 
dates were obtained, the seaward decrease in ridge 
elevation could be attributed to a rclatov fall m sea- 
k'vvl of - I m, However ihe evidence is inconclusive 
without subsurface stratigraphy and dotes; all ridge 



fig. 1. Location map showing portion of the South Australian coast and iocation of the 20 sites (1-20) discussed 
m text. See Tabic I for names of sites. Lower insert is of Western Cove (sites 15-19) on Kangaroo Island. 

crests arc also within the reach of contemporary 
.storm swash, hollowing progradation the ridges 
were buried by dune transgression, and latei 
exhumed as a deflation surface, suggesting thai 
shingle ridge formation preceded the deposition of 
barrier sands, corresponding to the finding of Burne 
(1982) ill noilhein Spencer Gulf. 

Iipson Cove — Shitif>/e Ridge 

Shingle ridges back many oi the sandy beaches 
along the western Spencer Gulf, particularly in the 
vicinity of the Tertiary conglomerates. At Lipson 
Cove (site 4, Fig. J ) a single shingle ridge (Fig. 3d) 
has been emplaeed landward of the present beach. 


Tabic I. Site hac&ttOn* and Chqwci&ri&tici 

H islam 


Marine Pncriw 




1579 km 

8 beach rid^Cs 


Point Germein 


7 beach t'oredune ridge: 


The Knob 


11 shingle ridges 


Lip^on Cove 


1 shingle ridafi 


lulka Well 


low beach ridges 

somhcrlv -wind waves 
southerly wind waves 
southed} wind waves 
>ou(herly wind waves 
northerly wind wave;* 


a- C OlTin Bay 

B; Morgan's I atuliny; 20tfy 

7. 5aji V>;ltei]Hik 

S, Yati£ie Bay 

9, Mount Dm ion Bay 

b. Western E\re Peninsula 

10 Peduhie Beach (^ 

11. Acramau Creek 

12. F.vre Island 

13. Laura Bay 
|4, Tbumlle B;tv 




4 be:u*U rsji^^s 
7 lutcduue tidies 
tidal Hals 
3 heath i Hives 

beach loredune ndge^ 
multiple beach-nd^es - spit 
multiple beach-rid^es - spit 
6 beach ridges 
3 Liiettiets -<ind mangroves 

ocens\ swell and N, wind svaves 
alia; pieviottslv low swell 
no waves 
low S wind waves only 

W wind waves 

W & S wind waves oceas, *vvell 
oceas. swell & W wind waves 
occas. swell & S wind waves 
occas. SifctfM 


15 Biownlow 2f> km 
lft. Brownlow (nolf course) 38 

17. Cvgnet Rtvei ?l 

IK. Mormon Beach 42 

19. Monison Beach 43 

20. Peimtiuuon Bay 156 

5 beach ridges 

7 beach lid^ei. 

2 chenier beach ridges 

hdaJ flat 

back barrier depression 

shore platform 

N & F wind waves 
N & E wind waves 
N & E wind waves 
N & E wind witves 
N &. E wind waves 
open swell 

'Mainland distance based on (SIRO Australian Coastal Invenrory knis km al SA' Vie border with km increasing 

to west to 32^7 km at SA/WA border ((ialknvav vt tfl \W4). 

'Distance on Kanproo Island bayed on clockwise measurement from North Cape (Shan & LVuhcinw.ham I9K6). 

(he location oftbe rid^ie su^e^ deposition prior 
to sandy beach formation, while its crest elevation 
of 4 m above ISLW is within the reach of 
contemporary sturm waves. 

hitka Well — Beach Rid&es 

Lulka Well (site 5, h'ijj. I) is a protected low wave 
encrpy site receiving only wind waves generated 
across Porl Lincoln proper by north-east winds. The 
small, isolated beach ridge plain consists of several 
low ridges which have prograded approximately 1 50 
m into the bay. Dates obtained from three holes due. 
in Ice of the innermost ridge indicate that this was 
tidal flat hv at least 3510 "•_ 80 vr BP (sample CS 
53! Table 2, Fie,. 3c). The dates (WK 837 ami CS 
Sit J indicaie upward accretion of 25 cm over 0.6 
ka, a rate of 0.04 mm yt : . hollowing the upward 
accietton of the lidal flats, some time subsequent 
to 2.5 ka BR. the low ridges prograded across the 
Hals 15(1 m into the bay. The inner samphire — ndal 
Hal tidal flat limit could be interpreted as tying up 
to 40 cm above the present lidal flat limi! suggesting 

a .slight fall tn sea-level. While De Deckker et al 
(19K2) also invoked a progressive tall in sca-kvel over 
the last 5-6 ka to explain decreased salinity in 
nearby Pillie Cake, the present data were 

The Great Australian Bight 
(Western Kyre Peninsula) 

The Great Australian Bi^ht (the Bight) is exposed 
to some of the world's highest and most persistent 
waves (Chellon el ul. 1 L )8<». However shelf 
attenuation fProvis &. Sieedman I9K5), coast 
oriental ion and the presence ol extensive nearshore 
calcarentte suhslantially reduces wave energy at 
many siies. evert on the open coast (Short el al. 
19Sfi). \i a result low energy beach tidges and 
elieniers are found in a number o( loeatioas 
particularly m the larger bays — Coffin, Venus, 
Baird ? Streaky, Smoky, Decres and Iburville (Fie. 
I). Nine sites were investigated. They are located in 
Colfin (4> T Streaky (2) f Smoky (1). Decres (1) and 
Tourville (I) hays, (Table K Figs 1, 4). 



Coffin Bay 

Morgan's Landing. 

Morgan's Landing (site 6, Fig. 1) contains four 
beach ridges backed by a back barrier depression 
dominated by a Melaleuca swamp (Fig. 4a). In 
addition, shell-rich beaehrock is exposed on the 
beach in front of the ridges (Table 3). Three dates 
were obtained. The oldest (WK 829), from [he back- 
barrier swamp, dates at 3690 ± 190 yr BP, with the 
rear of the innermost ridge dating at 2540 ± 60 
yr BP (WK 827) and shell in the beach calcarcnitc 
at 2680 ± 80 yr BP (CS 527). This suggests that 
progradation of the three ridges occurred around 
3-2.5 ka BP and that since then the shoreline has 
been relatively stable. This stability is interrupted 
by west to east moving pulses of sand (sandwaves) 
that alternately prograde (such as when beach 
calcarenite was deposited) and erode the beach 
(such as at present) as they pass. 

Salt Walterhole. 

A I km wide series of at least seven low foredunc 
ridges has prograded north-eastward of the Salt 
Waterhole (site 7, Fig. 1; Fig. 4b). The ridges 
apparently prograded at a time when a tidal channel 
existed between the Waterhole and the present Point 
Fongnose. The eastward growth oi' Point Longnose 
subsequently closed the channel and terminated 
ridge progradation. Fhiee dates were obtained from 

the back barrier depression and the swash deposited 
base of the first (oldest) and fourth ridge. The back- 
barrier sample (WK 829) dated at 3440 ± 60 yr BP, 
the inner ridge (WK 830) at 3930 ± 60 yr BP, and 
the fourth ridge (CS 528) at 3190 t 80 yr BP. These 
dates suggest a period of ridge progradation 
heginn'mg about 4.0 Ka and continuing to some- 
time beyond 3.2 ka BP, probably around 3.0 ka BP. 
At this time it is likely the channel was closed by 
the eastward growth of Point Longnose causing the 
ridges to be cut off from periodic ocean wave attack 
and sediment sources. 

Yangie Bay. 

Yangie Bay (site 8, Fig. 1, Fig. 4c) is a subsidiary 
embayment in the southern part of Coffin Bay. It 
is a zero wave energy environment where Holocene 
sedimentation is dominated by tidal and biogenic 
activity. One date (CS 529) was obtained fronuidal 
flats surrounding an outcrop of Pleistocene dune 
calcarenite and should indicate the commencement 
of tidal flat progradation in this part of the bay. 
it dated at 2780 ± 80 yr BP (Fig. 4d). The other 
date (WK 832) was taken at 60 cm depth beneath 
samphire flats extending from the calcarenite and 
dated at 2080 ± 60 yr BP. Both dates suggest that 
substantial tidal flat progradation did not 
commence until around 3.0 ka BP. Stratigraphic 
examination of the site (Fig. 4d) also suggests that 
there has been little if any change in sea-level .since 
that date. 

Fig. 2. The beach ridge plain near Glensea is located amongst longitudinal Pleistocene dunes (D). The transet (Fit>. 
3) extends from A across the ridges to B on the inner iniertidal sand flats. 





i 2 i ; 


Site 1 'Glensea' beach ridees 

4 5 5 

•A A r — \ y\ 7 


2S60 330 

i i i 

CS52-5 \/ \ / 

4 \>r CSSioN. Tidal Flats - 


1 1 1 

Site 2: Pc 

)tru Cermetn 

/I / I Scarp 

Site I: The knob 



d 6 A 
1 /V S\l\ 

A 4 

■ \ i i n 


/ \ 2 / \s 

3220 '80 V 

V \ *' 



V T ' \ 

1 1 





I \ I 

Site 4: 1 .ipsom Cove 

Site r > rulka Wells 


\ A Shingle 



\ Sand 








ry~~ ^*~ > > N Samphire Facies 

a WK837 29Q0+60 *"Vf" 


b.CS53t 35101S0 
C.WK836 2420 ^60 




1 1 1 I 






DISTANCE (meters) 
Fig. 3. Spencer Gulf Held sites, Chronological ridge numbers indicated above some ridges. 

Mount Dutton Bay. 

On the north-western side of Mount Dutton Bay 
(site 9, Fig. 1) is a number of very low energy low 
beach ridge sites. One of these was surveyed and 
sectioned (Fig. 4e). The innermost sample (WK 833) 
dated at > 40,000 yr BP indicating Pleistocene 
material. A sample obtained from tidal flat deposits 
below the base of the innermost ridge yielded a date 
o\ 6330 ± 90 yr BP (CS 530), the oldest Holocene 
date recorded in this study. This date confirms the 
arrival o( the Holocene sea-level stillstand by at least 
6.3 ka BP, in agreement with its arrival in Spencer 
Gulf (Bclperio et at. 1984) and Australia in general 
(Hopley & Thorn 1983). This date also is supported 
by the age of the first swale which dated at 4590 
± 80 yr BP, the third oldest date in this study. The 
elevation of the samphire — beach ridge, and tidal 
flat — samphire contact between ridge 1 and 2 (Fig. 
4c) suggests a slight ( - 50 cm) fall in relative sea- 
leveL The timing of this apparent fall however 
cannot be determinded from the existing data. 

Streaky Bay 

Perlubie Beach (south) 

Perlubie Beach (site 10, Fig. I) is located in south- 
east Streaky Bay and consists of a 1 km wide plain 
of Holocene beach and foredune ridges (Fig. 5a), 
A single date was obtained from the shell-rich swash 
/one immediately behind the innermost Holocene 
ridge (CS 499). It dated at 3290 ± 90 yr BP, again 
suggesting a time gap between the stillstand and the 
onset Of low energy beach ridge progradation. 

Acraman Creek 

The Acraman Creek (site U, Fig. 1) region in 
northern Streaky Bay consists of a 10 km section 
of shoreline dominated by multiple episodes of 
recurved spit development and bay ward beach ridge 
progradation (Fig. 5b). The shoreline at its 
maximum has prograded up to 3 km seaward. One 
date was obtained from the inner portion of the 
back barrier depression. Whole shell samples (CS 



TABLE 2. Hadtocarhon Dates 

3, Eyre 





C'SIRO h'nvirunmeni 


(in MSLW ) 

Age I 
Yrs BP 

r 1 o 

Yrs HP 


L 1 O 







CS 510 

CS >ll 

CS 524 
CS 525 
Wk' 826 




Back Swale 
Beacn Ridge 
Beach Kidgc 
Beach Ridge 






25 SO 
25 30 








CS 526 

Poim Genuein 


Beach Ridge 











WK 837 
WK S36 

Tulka Well 
Tulka Well 
Tullca Well 



Tidal Flat* 
Tidal Flats 
Tidal Flai.s 










Coffin 1! 



WK 828 

WK 827 
CS 527 

Gate Wacerhcrte 
Morgans Landing 
Morgans Landing 


Inter Bairiei 
Back Swamp 
Beach Calcarenitc 




inler liclul 

4| "0 










Wk 830 
CS 52X 

Sail Waierhole- 
(Coffin Buy» 
(Collin Ba>) 



Back Swamp 
Swash Deposit 
Swash Dcposh 



- 1 





3 190 




CS 529 
WK 832 

Yaugie Island 
VampE Island 


Inter Tidal 
Inter Tidal 











WK S33 
CS 530 

WK 83-1 

Mt Dutlntt Hay 
VI i Duo on Bay 
Ml Otiilon Bay 



Inter Tidal 
(met Tidal 
liner Tidal 




6790 * 90 
S050 i 80 

> 40.000 
6330 i 90 
1590 < NO 

Great Australian Bin 



CS 499 

I'crlnbic (south) 


Inaei iic'iuh Ridre 









cs m 

Acantmnn Creek 


1 liner mo.M Swale 









CS 469 

Poinl Collmson 


CS 169 

Poinl Collimon 


CS 509 

Evie Island 

3 71 

CS 503 

Evrc Uland 


CS 506 

Ryre Island 


CS M)5 

Pyre Island 


WK S15 

Laura Bav 


CS 504 

Laura Bqj 


WK 824 

Laura Bay 


CS 502 

1 aura Buv 


C.S 501 

Laura Ba\ 


WK 825 

Laura Bav 


CS 500 

Lnura Bay 


WK H16 

rourviJle Bav 

3-3-1 Wk 817 

1-*? WK 8IK 
3-36 W K 819 
3-37 WK 820 

3-38 WK 821 
b, Kangaroo Island 

Davenport Creek 
Duvenporl Creek 
Davenpon Creek 
DmenpOM Creek 
Davcnporl Creek 

261! Shell Rock 

261 1 Shell Rock 

2635 lmei Tidal 

2635 line- lidal 

2635 Inler lidal 

2635 [tiler tidal 



Beach Ridge 

Beach Ridge 

Beach Ridge 

Beach Ridge 

Beach Ridge 

Beach Ridge 

39 Beach Ridge 

Chenier Ridge 
Chenier Base 
Chenier Base 
Chenier Base 








Kl 13 WK K39 Mumson Beach 
KliO Wk 842 Cyenel River 
Kl M Wk 841 CviiKM River 


Reach Kitlge Swale 
Shell Ridee 
Shell Ridge 





- I 





2 5 






1 - huIk;j:^n, iiri'-iirvevfii, csrnvie only 

2 Ctareiajoti I aclai or 460 Uaif^ ihcd, 









1860 ± 


1400 i 


3300 - 


2840 . 


3680 2 


3220 [ 


377[| * 


HI0 v 


6140 • 


56M0 rt 


2740 . 


2280 t 


2230 . 


I77fj * 


2520 f 


2060 | 


2760 + 


2300 ± 


34SO 2 


3020 h 




398(f t 



2470 + 


201O j 


2R60 . 


2860 1 


2570 i 


2S7t) - 


25 40 - 


2540 -f 


2690 s 


2690 t 


5040 - 


4580 - 


3: io 


7650 • 




2290 * 


Ho-,wii;tn lv lluivev flQKJj: 


A. I). SHOOT. R. C. BUCK1 UY £ D. G 1 aillbHINGM W 

5931 gave a dale i»l SSSO • 90 yr BP the second 
ulticRl Hi the Holocene daiev 'I hiv also support* 
the minimum n^c ol ihe Holocene stillsumd in i his 
jeyiuo at >& ka BP 

Lwe Island (sue 12. Hp- 1). winch partially blocks 
Smoky Hay is die only unconsolidated island in The 
Bmhl. It consists of a multiple series of onlapping 
recurved spits, bench ridees and low foredunes 
which surround a core ol samphire -- mangrove 
Hals 0"ig. 6). A nansect was made across Ihe 
westernmost and oldest ol' the existing ridges (Flp. 
5c). The date obtained from the three swales and 
behind Ihe present beach indicate that this scciion 
of the inland began building aromul 3.8 ka BP ami 
progruded to around 1,4 ka BP, This section is 
presently eroding as sediments are rewoikcd 
eastward along both sides ol the island to pio^radc 
the recurved spit complexes. As ihe island may have 
been reworked eastward piioi to 4 ka BP |N dates 
can only be used as a guide to the age of the oldest 
section ol the existing island, and do not necessarily 
indieale (he commencement at island formation 

Laura Hay 

Laura Bay (site I3 t Fig. I» is a 2 Km wide Day 
within Deeres Bav. Jt contains n 1.2 km wide heach 
ridge plain fronted by several hundred metre wide 
ialcrtidal sand flats, The bay receives essentially no 
ocean wave energy as attested bv the growth of 
maugitwes on irs northern sandllats. A sencs of 
seven dales obtained from two swales and the base 
of live ridges (Fig. 5d) .suggests that progradation 
commenced alter 4.0 ka BP (CS 500). A period ol 
sJow progradation (100 m between 4-3 ka BP, ridges 
I and 2), was followed by rapid progradation (500 
m> between 3-2 ka BP (Fig, Sdji Ml ridges are 
wahm the reach of present storm swash elevations 
and do not suggest any appaieul change m sea-level. 
Like the Glensea ridges, the 2.3 ka age for the outer 
active t idge (6) may indicate age contamination by 
older shells, and hence a yuuugei age may be 

lounillc Bay 

Tuiirville lia\ (Site 14, Eft, I) is the westernmost 
enclosed bay in S. Ausi. and contains rlie 
wcsicrnmost mangrove woodland in Souih and 
southern Australia. A total o\' five daic* wcic 
obtained in the bay, lour I'rorn Davcnpot t C reek and 
one from the northern side ol ihe has (Fig. 5c), 
Davenport deck consists of a 10 km mangiove 
Woodland described by Bullet ci af. (I'>T7k willun 
which he rluce discontinuous chcuieis(tr:i ( ?). The 

second of the three vhenicrs was sectioned and 
provided ;i sequence ol llirec date.-. (Wh §20', SIN, 
819, Fig. 5^, These dates confirm ihe emplacement 
of l his second chenier at between 2.5 and 2.0 ka 
BP, and the development of the underlying lldal 
flats bv 2.5 ka BP. Mangrove stumps on theruuuh 
side of Davenport Occk, presently being huoed by 
dune transgression dated at 2690 i 50 (\\ K K21), 
comenipoianous with the chenier base mangroves. 
The dale on ihe noithetn shore of loinvitle Bay *'as 
obtained from the intcnidal base ol the innernu)sl 
beach i tdgc (Wk 81bL Its date o\' 2190 * (>0 yr BP 
coincides with the development ol Ihe second 
chenier. these dates are confirmed by Ihe rvMills 
of Bclpcno ft a/. (I9HK) who present ihe result*, of 
13 vibroeoj'cs across Tourville Ray. TIk> tonm! the 
basal scagiavs lacrcs beuan accumulating 
about f>K0(> yr UP, wilh ihe embaMiicni laigclv 
infilled by 5000 yr BP Beuch ridge formation 
followed the mainline ol ihe scagtass banks bv 
inlcrtidal and mangrove lacics. Their innermost and 
single beach rrdg* date of IH40 yr BP is in 
agreement with our findings. 

K;m! ;-■;[.. j 1 1 Island 

Most of the Kangaioo Island woast is typified by 
IjTglj betltoek and ealcaicnite chl'ls exposed lo tu^li 
wave and wind conditions. However the nonh east 
leeward part o( che island contains rhree 
embavntcnis — the Bay ol Shoals, Western and 
Eastern Coves which offer varying degrees of low 
cuetjjy Holocene shoielmc piogiadalion. Western 
Cove has experienced rhe most substantial 
piogradation with sedtmem being delivered 
alongshore Irom its eastei n and western arm, as well 
as from the tymnet River and ffe tiftt deposits. Lour 
Mies were examined around the shoielmc (bigs 1, 8). 


A series ol* I'ive. low, shells, Mvash-deposiled 
beach ridgev wa< surveyed at Brow tow (site 15, 1'ig,. 
1, rt^i. Sa). Their uniform elevation suggests no 
apparent change in sea level. No dales have hen 

Bnnv/tlow jjoff counv 

Seven low shelly t'idfes were surveyed immediately 
soulh ol the Brownlow golf Ci>utse (site K». Hg. 1 1 
and again iheir relatively uniform elevation gives 
no indication of sea-level change (Pig. Sh) 

Cyi>ju'/ Rfvtv 

A series ot low. discouln'uious beach ridge and 
possibly cheniers were surveyed I km south of ihe 
Cvimct Kiver mouth (>iie IT, Fig, I), The two ridi'es 



Site 1 6: Morgans Landing 

Hulqcene Beach 
Ridges ^ 

Site 7: Salt Watorhole 

Coffin Bay 

Fig. 4. Coffin Bay field sites, Sites 6, 7 and 8 were not surveyed. Sketches based on 1:10,000 aenaJ photograph 



bite 10: 
Perlubie Beaeh<g 

Site 11: Acranifin Creek 

ti. CS503 5650 90 


A N 


*£/ X ^*^<Ev Acraman Ck 
Beach Ridge* -> ^^ )/ 

PL Lindsay 

Streaky Bay 


Site \2. tyro bland 

c. CS&05 3a to ieo 

d.CS506 3220 go 
e.CS508 2840 90 
F. CS509 MOO +80 


-i 1 — 1 g. r,5500 398Q19Q 

Site H: Laura Bay , cssoi 2300-90 

j CS502 2060 1 90 

k.WK624 1770 70 

Shellv S-and I, WK81& 5680 90 

P ,s/ m. CS504 2280 90 



200 400 600 600 10Q0 1200 1400 

Pig. 5. Western Eyre Peninsula field sites. Sites 10, 11 and 14 not surveyed. Sketches based on 1:10,000 aerial photographs. 
Site 12 itirvfy elevation* suspect, distances correct. Chronological heach r idye numbers given for sites 12 and U. 

yielded dates of 2650 . 60 and 2290 + 50 yr UP 
( WK 840, VVK 841). The ridges are now fronted by 
300 m ol densely vegetated samphire Hats (fig, 8c). 
1 heir distance from the present shoreline and the 
elevation of the samphire flats could have resulted 
I'mm a slight fall in sea-level subsequent to their 
formation. However this cannot be confirmed from 
the present data. 

\1om\on Beach 

Two sites were examined at Morrison Beach (sites 
IS and 19, Fig. 1). In a waterhole al (he eastern end 

of the beach (Site 18) (he elevation of imeitidal 
shinule exposed in the waterhole is - 0.5 ni higher 
than the present beach shingle fEig, 8d), and could 
be attributed to a slight fall in sea-level. However 
the elevation of the inner shingle lies within the 
reach of storm swash activity. 

The bcaclvforcdunc ridge plain at Morrison 
beach consists pf a 2-3 km wide plain that has 
ptograded northward and westward into the bay 
(Fig. \\ Morris 1976). Shells obtained from a swale 
behind one of the innermost ridges (silc 19) (WK 
339) dated at 45S0 + 70 yr BP, indicating that 



Fig. 6. A view looking eastward along the Smoky Bay side of Eyre Island. The subtidal sand-flats are backed by 
a low energy reflective beach, and multiple recurved spits/beach ridges with samphire flats and mangroves 
composing the core of the island. 

Fig. 7. A view southeast across Tourville Bay shows the Davenport Creek tidal channel and sandflats backed by mangrove 
woodlands and the discontinuous chenier ridges (C). Active dune transgression (D) in the south is burying the 



TABLE 3. Beach ridge molluscs in approximate order of abundance 

a, Interiidal mollusc species commonly found in Eyre Peninsula Holocene beach ridge and swale deposits. All these 
species are associated with low energy mud/sand flat situations, generally with associated Zostera beds or silted rocks 
etc . . . Coxiella sp. is usually associated with salt pans or salty lakes. Bembicium melanostoma and Salinator fragith 
are found on mud Hats at the upper intertidal zone in sheltered estuarine areas. Anadara trapezia is usually associated 
with Zostera beds; it is extinct in S. Aust. 

b, Rocky shoreline species {also found in Eyre Pensinula beach ridges). 

C, Kangaroo Island beach ridge molluscs, primarily in Western Cove region. 

Katelysk peroni (Lamarack) 

Katelysia scalarina (Lamarck) 

Amesodesma erycinaea (Lamarck) 

Amesodema cuneata (Lamarck) 

Spisula trigonella (Lamarck) 

Anadara trapezia (Deshayes) 

Brachidontes erosus (Lamarck) 

Batillarielta estuarina (Tate) 

Zeacumantus diemenensis (Quoy Gaimard) 

Salinator jragilis (Lamarck) 

Thalotia conica (Gray) 

Prothalotia sp. 

Bembicium meeanostoma (Gmelin) 

Coxiella sp. 

Phasianetta australia (Gmelin) 

"Bulla fco/flmctf" Hedley 

Diala sp. 

Tellina (Macomona) deltoidalis (Lamarck) 
Trichomya hirsuta (Lamarck) 
Cominella eburnea (Reeve) 
Austracocholea constricta (Lamarck) 
Phasianotrochus rutilus (A. Adams) 
Sanguinolaria biradiata (Wood) 
Mactra Pura (Deshayes) 
Futvta tenuicostata (Lamarck) 
Ostrea angasi (Sowerby) 
Bembicium cf. auratum (Quoy & Gaimard) 

Katelysia scalarina (Lamarck) 
Katelysia peroni (Lamarck) 
Katelysia rhytophora (Lamarck) 
Zeacumanties diemenensis (Q & G) 
Mesodesma cuneata (Lamarck) 
Fulvia tenuicostata (Lamarck) 
Austrocochlea concamerata (Wood) 
Nassanu pauperata (Lamarck) 
Pisulatrigonella (Lamarck) 
Batittaria estuarina (Tate) 
Nerita atramentose (Reeve) 
Brachydontes erosus (Lamarck) 
Tellina (Macomona) deltoidalis (Lamarck) 
Cominella lineolata (Lamarck) 
Cominella eburnea (Reeve) 
Austrocochlea concamerata (Wood) 
Salinator sp. 
Diala sp. 
Bittium sp. 

initiation of this substantial and continuing 
shoreline progradation commenced by 4.5 ka BP. 

Shore Platform Evidence 

Shore platforms cut into Pleistocene beach and 
dune calcarenite are widely distributed around the 
western Eyre Peninsula and Kangaroo Island open 
coast. Whilst shore platforms are highly 
questionable indicators of sea-level (Twidale et ai 
1977; Hopley & Thorn 1983) they do warrant brief 
attention in regard to possible changes in Holocene 
sea-level in the Bight and Kangaroo Island. 

Three platforms in the Bight were examined for 
evidence of Pleistocene sea-level (Buckley et ai 
1987). Two of these platforms contained a well 
defined upper, inactive Pleistocene platform, 
fronted by a wide active intertidal Holocene 
platform (Merdayerrah and Clare Bay; Fig. 1). 
Subsequent low altitude aerial reconnaissance of the 
entire Eyre Peninsula and Bight by ADS and DGF 
revealed the existence of relict elevated platforms, 
presumably of Pleistocene age along much oi the 
calcarenite coast. Active horizontal shore platforms 
occupy 440 km of the Eyre Peninsula coast (Short 
et ai 1986) and are usually located intertidally close 
to mean sea-level. All western Eyre Peninsula 
platforms examined possessed a single active 

platform surface. On Kangaroo Island, however, 
Short & Fotheringham (1986) describe a Pleistocene 
platform at Pennington Bay (site 20, Fig. 1 fronted 
by two Holocene platforms at different elevations 
(Fig. 8e). The upper platform (No. 1, Fig. 8e) lies 
50 cm above the lower platform (No. 2). The 
difference in elevation and abrupt drop from 
platform 1 to 2 may be a result of a slight fall in 
sea-level ( - 50 cm) producing a lowering of the level 
of saturation and hence platform erosion surface. 


These data have implications both for the 
evolutionary history of the low energy beach ridge 
sequences and for mid-late Holocene sea-level 
history in the Bight and on Kangaroo Island. 

Beach Ridge Progradation 

The initiation of beach ridge progradation at nine 
of the 12 sites dated did not commence until 4 ka 
BP and in some areas as late as 2 ka BP (Fig. 9). 
Most sites indicate an hiatus between the Holocene 
sea-level stillstand and inner ridge building of up 
to 4 ka. This is in general agreement with Burne 
(1982) who found that in northern Spencer Gulf the 






Site 15: Brownlow 

AeoMan Sand 


Shoffy Sand 



§ 3 


>- 2 


Site 16; Brownlow (Golf Course] 





Site ]/ : Cygnet River 

a WK840 2650 GO 
b.WK841 2290 ' 50 

Fine Sand 



T7T7T77 T777777T7 



it i 'lite \\\\ Mornson Beach 




Shore Platform 




6 I Site 20: Pennington Bay 





60 10 

DISTANCE (meters) 
Fig 8. Kangaroo Island field sites. Beach ridscs are numbered chronologically. 



development of low energy beach ridge plains 
awaited sufficient subtidal carbonate production 
and as a result progradation did not commence until 
4-3 ka BP. Belperio ct al. (1 988) also concluded that 
carbonate secreting organisms aggrade and 
ptograde the subtidal sand hanks in association 
with the subriUal seagrass facies. Once (he banks 
aeeumulaielo low water level, subsequent intertidal 
and beach ridge progradation can occur. They 

found I his sequence a! Tourvillc Bay on Tyre 
Peninsula as well as several sites in Spencer and St 
Vincent Gulfs.. 

In Gulf St Vincent, Bowman & Harvey (1986) 
found that the extensive Le Fevre Peninsula beach 
ridges have been prograding continuously over the 
past 7000 years with pluses in ridge formation 
apparently linked to longshore (northward) 
sediment supply. This higher energy west facing 



system (modal Hb = O.S m't represents a transition 
between the low ciiergv beach ridges discussed in 
this paper and the litlgh encrgv open coast regressive 
dcposiis winch experienced rapid progradatum 
following ihe saliscand Their mode and ivquence 
of evolution of the Peninsula is Therefore not 
directly fippliUHfels to 'be lower energy systems tit 
nil tfh&h experienced a substantial lag in ridge 

On the two sites where a sei|uener t*1 beach ridges 
*$S dated (excluding, tyre Island) The vouugesi 
ridge* dated 20)0 yr BP (GlenseaUnd I770 vr BP 
< I aura Bay), suggesting thai ridges ate either new 
currently progruding or older material is being 
deposited during ridge building. Whilst Tidge 
cessation is ui agreement with sedimentation models 
in higher energy sites in south-east Australia (Thorn 
el dl. F98I), it is surprising in an environment where 
moM sediments aix* deiived from in y/frV carbonate 
production from fronting iniei tidal and subtidnl 
sand Hats, ihe so culled Vuibonatc factory' | But lie 
1912). In northern Spencer dull', Burnc found 
periods of shoreline (ie. sandflat) progradntiun, 
ahei nature with beach-ridge piogradation. The 
mnsi notable shoreline prog r<*)dai ion WHS between 
3 and 2 ka BR On ihe »>re Peninsula, hcAvexer, I hat 
was the period of maximum beach ndge 
construction. A similar contrast applied to the 
period from 2 ka BP to proem; beach ridges in Ihe 
norih-eaM Spencer Gulf prograded, while iliose in 
the Pyre appear to have remained static. Evidently 
nioie information is required before genetal models 
..I *s Aust. bench ridgc-sandflat progrudalion can 
be clarified. 

tyre Island has a slightly younger oilier ridge 
U40G yr HP) and the island is continuing to develop 
new recurved spits, However this may be accounted 
for b\ erosion and dowudrift reworking n( island 
<edimeuts imbcr than external sand supply. 

Htfttfcette Sea-Level 

The exaci liming of the Holocelic SCtt-locI 
maximum cannot he resolved from these dales 
Howos'i the date of 6330 • W yr BP from Little 
Douglas as well M \\k Sfi5Q i 90 yi date Irom 
Acraman Creek lend support to the general curu- 
adopted by Bclperio et oi (W84> (uj Spencer Gull 
and I lopley & Thorn <l¥83) for Australia a>. a whole, 
Thai is it appears lhat ihe scra-lcvcl reached it\ 
present position M Wj fo HP In the Great 
Australian Rigid. 

barren, I K [\%H) I Witt QuaiCMHtr? evapotaic 
taint male deposit innal citviitifimwu* til Nuwli An .mtli.t. 

PhO dirw, H.mlcts Ultlv, Ai'MHidc (Ulipubl I 

In evaluating evidence Col relative sea-level history 
during (he mid to late I lolocenc, tectonic differences 
between the ihree main regions, the Bight, Spencer 
Gulf and Kangaroo Island must be laken inro 
account. The most stable region appears lo be 
western liyre Peninsula (the- Bight) Tectonic stability 
is suggested bv the widespread occurrence of the 
-j 2-3 Pleistocene shore platforms (Buckley et qL 
1986), the continuity Of the presently active bhorc 
pluifouns. the single intertidal solution notches in 
limestone cliffs in Coffin Ray and the c\ idencc horn 
dated beaeh ridge and tidal sites (Hgs 4. 5), In Ihe 
Bight the sea-level maximum was reached at least 
by 5650 yrs BP (Fig, 5-l» and has been stable since. 
In northern Spencer Gulf there is, however, evidence 
for Holocene- tcctonism. Belpeno et (ti (1984) 
proposed n sea-levrl curve foi northern Spencer 
Golf which has a slillstand being reached at -6,5 
ka followed by an abrupt fall to present sea-Ievet 
al 1.7-1,6 ka BP. The) attribute Ifcfi relative fall to 
local tectonism. The preseni SptiilWl Gull dala are 
not sufficient lo add to the Belpeno et ol. cut ve and 
may well be lecionicalK or isostaticnlly distmel. The 
Glensen beach ridges and swales slope seaward I Fig. 
,3a); however this is not conclusive evidence lor a 
shphl lalt in sea-level without subsurface 
morphostratigraphy. h could atst> represent ii 
decrease in wave energy nsihc I km wide intenidal 
and subiidal sand flats aceunuilaicd, I .ikewisc, while 
othei Spencer Gullsitc-v logeiher with Vailgie Island 
and Mount Duiton Bay sites may suggest a modest 
fall in sea-level (< I m), criocs inherent in sut veying, 
facies ideniiftcation and elevation, dating and 
establishing reliable datums. place all of the possible 
change^ uithin the sampling error bands. C ouse 
viuently there is no conclusive evidence lui a small 
relative la!l in sea-level II any Itas occurred it would 
be only on the order of a few decimeter Together 
Willi the Bight data this suggests a relaiivels stable 
sea-lesel since the sea-level maxima m or prioi 10 
6300 yr BP 

Summar) and Oinclnsiims 

1 "he following scciiicnce oi event> appears to htiu- 
occiu red along the low energy Bight, Evrc PdlUWll H 
and Knngaioo Istantl coasls .since the llolitceite sCil 
level siilKiaud 

7-6 ka BP The stiMsland was achieved by at least 
6331) t V0 yrs. BP and most likely by u.G-6,4 fa) 
RP based on evidence from northern Spencet Gulf 
(Bclperio <Y Ol 1984), 

6-4 kn BP Following rhestillstand most low energy 
sections <>\ ^-oast experienced mininuil shoteline 
progrddalion. E\cepOous were arras wheie shinel^ 
Wcis iniinediatelv Available, part icn lot ly mSpenccr 








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A. I). MtORI. R. C fJlK'Kl. I Y & D O. I OTHKKIlNt .MAM 

Gut!. In these areas siorni shingle rjfclgtta were 
developed lolloping (he siillstand. Whilst no dales 
arc available from ihc ptesem data, daiin^ ol similar 
ridecs in northern Speneci Gulf by Bume (}9&2) 
toond proetadaiion commenced mound 6 ka BP. 
Only thrOC heach rid^c siics (Mount Dutton Hay, 
Acraman Creek and Morrison Beach) have hack 
barrier deposits older llian 4 ka BP No beach ridges 
were dated prior to 4 kn HI*. However prior m beach 
jtfdge foimatiou the substantial intei- to siib-iidnl 
Atfld 1'kus would have to he building during this 

4-2 ka BP. This was apparently a period of major 
beach lidjic and limited chenier development in 
Spencer Gull, the ttk'hl and on Kangaroo Island. 
Thecuuinieiiecmcn! v( lidge construction has been 
dated as early as .WO i 60 yr (WK 830) iti Col Tin 
Bay tl-iy. 4b) and as lute as 2I90 ± 60 (Wk SI6) 
in Tourville Bay (Tig.. 5c). The mosi logical 
explanation of this svireo in ridjie lormalion i* the 
development ol I he inlet- lo sub-tidal carbonate 
platforms' and sediments required |o both support 
and supply sediment lor i ide.e development possibK 
coupled with a slight fait in sea-level. 

2-0 ka 111 1 hi^i. y is noi repicsentanvc tit' low energy 

deposits as many of the samples were selected to 

date the ulcteM ridycsvsalc. All that can be said is 

lhal on i he two sequences dalcd, the vouuyest tiflftC 

dated at 2010 and 1 770 vr BP ( Tigs 3a, 5vi), 1 (owever 

ihete is abundani evidence ol eoiitctnpotat'v ridge 

formation, particularly in areas oi i-ecurved spits 

(ee. I'.vte Kland, Aciaman Cieek, St Peter Island). 

Possible explanaiions for the outer ridge is that in 

areas u! shoie normal sediment supply there is a 

preseiu hiatus in ridge formation, tu possibly otdet 

shells aie beine ineocpotated in outer xkigc deposits 

io j'ive an anornolous awe. Whalevei the cause ihesc 

systems require further investigation In conclusion: 

I I ovv energy beach ridges and mmiic ehenieis 

occupv fti km (4 ,, 'i0 o( (he western Kpeneet Gulf 

and Hijiht, pailkularly in the larger more 

shellered hays, Thev also occupy 14 km of coast 

in ihc noith-cavi of Kangaroo Island, Most 

sequences consist ol 2-10 riciees (mean 4*5 in 

Spencer Gull) 

2. I he rtd^cs aiv composed ol predominantly 

carbonate shell dclnlns liable 3) apparently 

sourced and lewoiked from the adpeen: iiacr- 
and sub-iidal sandl'luts and udjaceiii joekv 
}, Ridw.e formalion awaiied the eonstrucooi ol \'i\v 
underlving inter- and sub-tidal sandlku fhe 
sandllats vscie required bOtfl lo jifi-«ij.1c a 
platform for ridge butldiue, as well Us ■> uady 
supply of material The increasing pioituuuisC 
of carbonate sediments is refleeled id ilic 
gulfwanl increase in carbonate content of die 
Gleiisea ridges (Pig- 5a). 

4. Once lite inlcrtidul sandfku platforms wne 
constructed, ridge lormalion otleu was rapid In 
places several ridges scpanned by widci *»\(i|y,s 
were deposited m several liuudied year-, usually 
between 4-2 ka BP. However age conlamiua! ion 
oi oiner ridges by oldei shells may mask a 
younger age and hence longer period ol' lidrv 

5. At present most active ridge foinsaoon is 
occurring m aieas of longshore supply, (ouUmti 
in the consiruclion ol multiple icoiHMd spits 
Shore normal supply may beuefe'li^iblc icMtltmn 
in an apparent hiatus in ridge coustnMion oi 
conversel> older material is being inc ap,r;Ued 
to provide anomalous ayes. 

6. The sea-Icvcl stiUsiand occurred phot hi \\J ka 
BP and lias been reialively stable sokv; 
particularly in the Bight. There iv inconclusive 
evidence of a slight {<l nu fall in sea-level m 
Spencet Ciulf and B (0.5 m) fall on Kanvamn 
Uland in (he late Holoecnc. 


All licld vvoik on Kangaroo Island ami ittc I uc 
Peninsula wrfs suppoited by the Coastal Pluttu '" i n 
lioatel ihicuiuh the Coasial Maua.eetiieni Biauch, 
South Austiaitan Departmeni of i'nviroiuiicM' <V 
Planning, The I yte IVmnsula ladioearbtm dating 
was ptovided by rhe S A- Oept ol Mines Sl t.ntia\. 
Molluscs weic mdcutified by Phil * oh ■man. 
Australian Museum, heme*, weie dialled bv 
Jcancue Neeson and ihc inaniisciipt tvped bv 
laneire Brennan and Ka% losici. Many tfumk- to 

Iti in i;i". A. I',, tr< i ir--. x ', A.. { . ^S* I H.^M.mow- 
\\ \ i si i i j . \ . OVsN) ^VcliH)i;tlL-i^^ill(TMl) /oii.iiitni 
ami il'c i-Mttuiinii rj| HoKiccne litial .SL'qtk'nccs y\ 
Mvnlk'in Vuymlia ftp i"5-4 l )7. fn (' I ic Boer; ut 
>ii its) "IkIc* IntUici'.ctJ Sc-clintcmai > Lmaonincnis 
ikhI Kivh V* (I) hS ulvt fnhf C-Sii), 

, U\u ••■ .'. R ^ IiO'-iIn, V A. LW&Jl \ review 

vi lIoloiLiit ',uii levels in^ouih Ai:sio»hi i|itt 4 . ti\ 


II Hoplev. tr.d.) AuMinlUui <a le\.'K in Mv Mm KO0O 


wart n review (Jt'ah. Cup. .\ Mouoy.ui. <w 
GcwrapDs hnnvs ( ook Univ., TttWiivville. 

'. _ J . os P.. i \i ii. H. A. <m v .M I he 

jllJUu'tiipb) &\ ti>a--Uil carboiiiifc l>anks >mhI Huhn OAfr 
KCU Ivvt-ls ol ra>r;hern Spencer Oull'. Sovuli \asu.'t»;i 

ltt>\\\iv., O. k Hasm w. N t\9#\) K-nlmiail mi iI.imiv 



of marine shells in Sotn!. Australia. Ausi. Archaeoloqv 

17, 113-123. 

<v: _ _. 1986 ' •.viiiorphie evoluiion of a 

Hotocone beach-ridge -. .iiuolcx, Le Fevre Peninsula, 
South Australia. 7. CW»wV Sfeft 2, 345-362. 
BiAklL 1 ;. H, C. StlOKl, S l>. & FOTHERINGHAYI, D. G. 

(1987) I. .iio l-ietsioceiu mm 'level maxima in the Great 
Australian Highl Sear. J; IK. 46-89. 

BtiKNr, R V, UW) Ivd.Mi. ; t. II of Holoeene sea level 
and ci';isial progradari >fj, northeastern Spcnecr Gull', 
South Australia BMP Jr Ausi. Geo/. Geophv*. 7, 

Brn.LK, \. I.. Di pers, A M, MrKii!Li>, S. C, & 
Thom\,, D. P. (1977) Distribution and sediments of 
maiupnv fiekl> in South Australia. Trans. /?, Soc S. 
Aust, il>1. 35-44. 

CHtliTON, D IV, Hl'HM.V k. I j£ P,\RKf., M. E. (1981) 
Global satellite rneasuuMk-nis of water vapour, wind 
speed and wave height. SiJflire 294, 529-532 

D\IIY, B.. Mil. MS, A. R , IWtDAlE, C. R. & Bourm-, 
J. A. (1979) Geology aid i.vomorphologv pp 1-38. 7/? 
M, X lyler. C. R. Twidale. & J. K. Ling, (Eds.) "Natural 
Histo'v o\' kangnioo Maud". (R, Soc. S. Ausi., 

De Deckki.r, P., Bolld, F, & Burne, R. V. (J982) Pillie 
Lake, Eyre Peninsula, South Australia: Modern 
environment and biota, dolomite sedimentation, and 
Holoccnc history. Tnm>- R. Soc. S. Ausi. 106, 169-181. 

Gai.iowav, R. W„ STORY. R.. CflDPKR, R. & Yapp, G. A. 
(1984) Coastal lands oi Australia, CSfRO Div. water 
and Land Resources. Natural Resources Series i. 

HAILS. .1 R. & Gostin, \ A. (Eds.) 1984 The Spencer 
Gulf region. Marine Gat/, it, 111-424, 

Hcsr, P, A, 1984 The fe Uiitritffl of sand "beach ridges' 
and fortdunes. Scorch 15. 289-291. 

HopleY, D. &. Thom, B. G. (1983) Australia sea levels in 
the last 15,000 years: a review. In Hopley. D. (Ed.) 
Australian feu levels in the last 15,000 years: a review 
pp 3-36, Occas. Pap. 3, Monog. Ser. Dept Geography. 
James Cook Univ., Townsville. 

Morris. B. I (1976) Investigation ol he,i\/. mineral 
sands, Momson Beach, Kangaroo Island. Mutem/ 
Resources AVv. 145, 34-46. 

Shoki, A. I). HV8tfa) Response of a high through low 
energy sands vousi to sea level transgression and 
stillstand, South Australia. Geographical Review 78, 

. (198*'-* Modes of Holoeene coastal dune 
transgressi^'i m southern Australia. Sedimentary GtuL 
55. 121-14. 

& KOI MkiciiAM, D. G. (l986)Coasial motpho- 

dynamics and Holoeene evolution o( the Kangaroo 
Island mas,. South Australia, Coastal Studies Ij nit Tech. 
Repu 86/1, i Coastal Studies Unit Univ. Svdnev. Svdney). 

, . & BtrKiLV, R. C. (19X61 Coastal 

morphodynamics and Holoeene esolution of the Evre 
Peninsula oast, South Australia. Coastal Studies Unit 
lech. Rept, S6/2, (Coastal Studies Unit, Lniv Svdney, 

__ & Hi I, P. A. (1984) Beach and dune murpho 
dynamics etJ the South East coast of South Australia. 
Coastal Studies Unit Tech. Rept 84/1 (Coastal Studies 
Unit, Univcsity o^ Sydney, Sydney). 

TitOM. B. G ( . MOWMAN, C. M„ Cili.l ISPIE. K., | "J Mi'l i , 

R. & Bah ! Ml, M. (1981) Radiocarbon dating of 
Holoeene b-.Mch tidge sequences in southeast Australia. 
Monog. 11, iX'pi Geography. R.M.C., Duntroon. ACT. 
IwiUALt. C R_. Bt«!,RNP T J. A. & Tvuuau, N. (I977| 
Shore platlV !<?, and sea-level changes in the t>ulh rciiion 
of South Athinflt? Trans. 7?. Soc S. Aust. 101, 63-74. 


by N. S. Pledge* 


A complete last upper molar (M ) and fragments of several other teeth found in a bore on the 
Talyawalka Anabranch, east of Menindee, N.S.W., are identified as the Pliocene species Euowenia 
grata De Vis. Euowenia grata is redescribed and refigured, and the new M is compared with the 
equivalent teeth of Diprotodon optatutn, Zygomaturuwss trilobus, Meniscolophus mawsoni, 
Nototherium mitchelli and Palorchestes azael. The Talyawalka specimen extends the range of 
Euowenia grata by almost 1000 km and suggests a Pliocene age for the deposit. 
KEY WORDS: diprotodontid marsupial, Euowenia, Diprotodon, Zygomaturus, Meniscolophus. 
Nototherium, Palorchestes, Talyawalka, Menindee, New South Wales, Pliocene. 

Transactions of the Royal Society of S. Aust., (1989), 113(3) 163-168. 


by N. S. Pledge* 


Pi EDGE, N. S. (1989) The occurrence o( a diprotodontid marsupial referable to Euowenia grata De Vis in 
western New South Wales. Trans. R. Sot\ S.Aust. 113(3), 163-168, 30 November, 1989. 

A complete last upper molar (M ') and fragments of several other teeth found in a bore on the Talyawalka 
Anabranch, east of Menindee, N.S.W., are identified as the Pliocene species Euowenia grata Oe Vis. Euowenia 
grata is redescribed and refigured, and the new M ' is compared with the equivalent teeth of Diprotocion 
optatutn, Zygotnaturuwss triiobus, Meniscolophus mawsoni, Nototherium mitchelii and Pa/orchestes azael. 
The Talyawalka specimen extends the range of Euowenia grata by almost 1000 km and suggests a Pliocene 
age for the deposit. 

Kr-v Words: diprotodontid marsupial, Euowenia, Diprotocion, Zygomaturus, Meniscolophus, Nototherium, 
Palorchestes, Talyawalka, Menindee, New South Wales, Pliocene. 


De Vis established the species Owen'ta grata (De 
Vis 1887), for a skull and jaws (Fig. 3, QM 1 519) 
found at Chinchilla, Queensland. Later he renamed 
it Euowenia (De Vis 1891) after finding the earlier 
name to be preoccupied by an annelid worm. At 
the same time, he named a second species, 
Euowenia robusta, for a set of dentaries found at 
Freestone Creek, near Warwick, Qld, in Pleistocene 
alluvium. Woods (1968) suggested that Euowenia 
rohusta is a junior synonym of Nototherium inertne 
which could effectively restrict Euowenia to 
specimens from the Chinchilla Sand, and to a 
Pliocene age (Woods I960; Bartholomai & Woods 

Euowenia grata occurs with certainty only at 
Chinchilla (Stirton et ai 1968). M. Archer has 
informed me (pers. eomm. 3.xi.l975): "In many 
cases, specimens should more honestly be referred 
to merely as undetermined small nototherines", and 
there is no evidence at present for any Pleistocene 
occurrence. Recently, Euowenia has been listed 
(Tedford, Williams & Wells 1986) in the Plio-PIeisto- 
cene Kanunka local fauna at Lake Kanunka, South 
Australia, but this record needs confirmation. 

The purpose of this note is to record a new 
occurrence of the species well beyond its previously 
known locality 

Fig. I. Locality map. While Water lake, lalyaw 
Anabranch, New South Wales. 


Material and Methods 

In 1969, Mr D. Walker of Talyawalka Station east 
of Menindee, New South Wales, presented to the 
South Australian Museum some unusual tooth 
fragments found in the sludge from a newly-drilled 
water bore in "White Water Lake" (32°25'S, 

South Australian Museum, North Terrace, Adelaide, S. 
Aust. 5000. 

143°18'E) on the Talyawalka Anabranch of the 
Darling River (Fig. 1). 

The new specimen is very fragmentary, 
comprising the complete left M , two enamel 
fragments with differing degrees of wear, believed 
to be anterobuccal corners of the protolophs of M ; 
and M 1 of the same tooth row, the anterobuccal 
corner of the metaloph of M 1 , the anterior half of 
M >, an unidentifiable molar fragment and a 
fragment of incisor. Because of the poor quality of 
De Vis' figure (De Vis 1887), the new specimen was 
for several years considered by the author to be a 
possible very large new species o\' Palorehestes, 
because it had a fairly distinct midlink between the 
lophs. However, direct comparison with a cast of 
the upper dentition of Euowenia grata (holotype 
QM F519) by courtesy of Dr M. Archer, has now 
proven its identity. The specimen also has been 









compared with teeth 01 caxts of Dipmtndon 
optanttn. ZygixtttaUirttt frifobiiSt Meniseo/ophus 

muwsotu, Xotaihenum mitehelli and Pahrehestes 

Material is from lite South Australian Museum. 
\delaide (SAM), Queensland Museum, Brisbane 
(OM) aral British Museflni (Natural History), 
London fBMNH). 

I he status of the genus i'nowemo is thought to 
he in do>,K; ,in<| 1P liccd 0\ Majtii revision (Archer 
iV Wade itm, hni until this h done it is fell that 
a i&tti i ort o! (he s|U- fc*. t r desirable, based 
partly on this new material. Tooth notation follows 
Archer (1978). 


Euowenia firuta De Vis 
FIGS 2-4. 

Owvniu grata De Vis, 1887, p. 100, jj|. 1-IV 

Upper premolar two-rooted, about half the length 
Of M : and triangular (De Vis 1887, 1888). 
Protoeone slightly smaller than parametacone. 
Protoeone and anterior part of parametacone 
joined by narrow transverse loph to form short side 
o\' a right triangle. Buccal side formed by buccal 
crest from parametacone to anterior end of the 

(ooth. Third side formed by anterolingual cingulum. 
Small deep central basin with rugose enamel. 

Upper molars large (Tables I, 2) and bilophodont, 
with lophs convex forwards and somewhat tumid 
with incipient midlink development — postprom 
crista posteromedially from protoeone and pre 
mctaeristat?) extending anteromedially from meta- 
cone. Midlink development virtually absent in M 
but increases slowly to its creates] eicul in M 
where pre- and post- cristas are conl'hu ni For l\ lew 
millimetres vertically. M ' suhreetu lyular; M 
trapezoidal, wiili mctuloph about fhiv;- -quarters 
width of protoloph. In M 4 and M\ mctaloph 
forms distinct dog-leg bend where midlink joins it. 
Anterior cingulum extends full width of tooth, bul 
is longer at its Ungual end. Posterior cingulum 
narrow, particularly on M where smalt pocket 
may be formed by weak postmetacrista. Only trace 
of a lingual cingulum may be seen, at bottom of 
transverse valley. Buccal cingulum better developed, 
across mouth of transverse valley. 

Lower molars more rectangularly elongate than 
uppers. Both protolophid and hypolophid arc 
oblique and concave anteriorly. Protoconid end ol 
protolophid enlarged by development of preproto- 
cristid extending anteromedially. Similarly, hypo- 
conid end of hypolophid is enlarged by cristid 
obliqua which forms posterior half of weak mid- 

TABLE I. Cheek Teeth Dimensions, Euowenia grata 

Except lor the premolar, measurements are made in planes parallel io and normal ro the lingual edge of the tooth. 

A-p I onjirh 
Am. Diarn. 
Pom. Oiam. 

SAM PI 4506 





QM 1-519 (epoxy east) 

SAM P14506 
l-M LM* 
















TABU: 2. Euowenia grata Dimensions as given by De Vis 0887} 

These are not directly comparable wfth those to Table 1, as M for sample is apparently measured alone rhc outer 


Mas. Breadth 

8 J 




15 JO 

14. s 


Pig, 1 Bkemm W*!q Pe Vis, I eft upper motai M o( SAM PI4506. Stefeopair photographs: pJOccluwit b) buccal; 
C) lingual view. Seale « I lid. 



FIG. 3. Euowenia grata De Vis. Holotype palate and dentaries. QM F519. Scale bar = 10 cm. 

link. In the holotype, lowest part of midlink is intact 
in last molar (De Vis 1887). Anterior cingulum 
almost full width of tooth. Postcingulum divided 
unequally into two parts at its highest point by fine 
submedian posthypocristid. 

Comparison with other species 

The Talyawalka specimen may be compared with 
this (Table 1) and other diprotodontoids. 

The upper molar M'is the same size as that of 
the holotype of Euowenia grata, and same order 
of size as Meniscolophus mawsoni. It is smaller 
than Zygomaturus trilobus and slightly more than 
half the length of Diprotodon optatum. 

The Talyawalka tooth differs from Diprotodon 
(SAM P23406, Lake Callabonna) in: its much 
smaller size, relatively higher crown, more convex 
protoloph, slightly narrower precingulum with 
greater development at the lingual end, a buccal 
cingulum across the end of the transverse valley, a 
narrow but strongly developed postcingulum, the 

absence of cementum, and particularly the presence 
of a Y-shaped hypoloph formed by a distinct 

It differs from Zygomaturus (SAM P29889, 
Naracoorte) in: its smaller size (about three-quarters 
the length), shorter precingulum, absolutely higher 
lophs, thicker lophs, presence of a distinct buccal 
cingulum across the transverse valley, deeper and 
more acute transverse valley, distinct premetacrista, 
more distinct postmetaloph-pocket, and presence 
of rugose/punctate enamel. 

It differs from Nototherium mitchelli (cast of 
BMNH M5002) in: its smaller size, its longer and 
better developed precingulum, presence of a buccal 
cingulum across the transverse valley, relatively 
higher lophs, presence of a premetacrista, and better 
developed postcingulum. 

It differs from Meniscolophus (SAM P13647, of 
which M' is unknown) in slightly smaller size, 
higher lophs, presence of distinct premetacrista, and 
narrow postcingulum. 



F% A, Cuowc/iia yjvta l)e Vis. CaM of left upper maxillary icmth row of holotype. QM f 519. 

It differs from the holotypc oi Luowenia gmfa 
(cast ol QM K5I9) in: having a distinct buccal 
clugulum with two small (one twice as big as the 
ol her | cusps across l he transverse valley, and a more 
distinct postmeialoph- pocket. 

II di Tiers from Palorchesfes uzael (QM F773, fide 
Oe Vis 1895) m: its larger size, relatively greater 
length, lack of a forclink, presence of a strong 
buccal cingulum across the transverse valley, strong 
postcingulutn, and absence of a complete rnidlink. 

The new specimens (SAM PI 4506) agree closely 
with the holotype, considering the difference in 
degree of wear and preservation In M\ the 
enamel is just breached so that a good indication 
of the height and lorm of (he tooth can be gained, 
The lophs arc high (equal to the width of the tooth) 
and stout so that the transverse valley is particularly 
deep and narrow, a narrowness accentuated by the 
bend caused by the premetacrista. Together, the 
mctaloph and piemelacrista have a distinct Y-lbrm. 
Contra Dft Vis (18K7), I found no indication of a 
forclink on Ihc upper molars of the holotype cast, 
but there is a slight protuberance in (his position 
on the new specimen of M '. 

The molar enamel is notably rugose and 
punctate, similar to thai of Dipmtodon and 
f : Mry:,yxoma (Archer 1977). In the worn M frag- 
ment, the anterior cingulum is heavily abraded by 
interdental appression and has a small cusp de- 
veloped at the anterolingual corner, The "rnidlink" 
is discontinuous and does not cross the transverse 
valley as an integral structuie. 

The incisor is too fragmentary to describe, save 
that it would be roughly circular in section with a 
diameter o\ at least 22 mm. Only a portion of the 
root ii preserved and, from Pe Vis' description 
(1887), it could be either upper or lower II, 


The teeth came from a depth of about 28 m, in 
a 1.5 m thick bed of coarse, yellow-brown dirty 
quartz sand, below sandy clay. The age is uncertain, 
but an early Pleistocene, or even Pliocene, age is 
possible, based on the absence of the species from 
laic Pleistocene surfieial deposits nearby — lunette 
sands dated at £5,300 years BP at Lake Menindec 
(Tedlord l%7) and 32,000 years old at I ake Mungo 
(Barbetti & Allen 1972). Although speculative, this 
determination is supported by ihe Pliocene age of 
the holotype (Woods I960). The specimen extends 
the known range of the lavon by almost 1000 km. 

It seems thai ihe bore drill struck a more-oMess 
complete skull and jaws of E'uuwenia because all 
fragments appear to relate to a single individual, 
(here being no duplication of elements. 


These fossil teeth, recovered from a wafer Km. 
on the Talyawalka Anabranch, east of Menindee, 
N.S.W., match those of Lttoweniu ?>rata (De Vis) 
and, because they are less worn than the holoiype 
add to the knowledge of the species. The occurrence 
extends the known range of the species. 


I thank Mr D. Walker lor clonal ing the specimen 
to the South Australian Museum, Mrs Debbie 
Brunker and Mrs Debbie Lowcry for typing the 
manuscript and Mrs I inda Btcsing for drafting the 


Ar< Hfr, M. (197?) Onnns unj Mibluinilial relationships 
ol Piprotodnn (Diprotodomidae, Marsuntelja), Mem, 
QUI. Sins. 18, 17-39 

(I97R) The nature of the molur-prcrnolar bouiutji v 

hi nnRutpnts and a rtantcrpreiation of the homology 
a\ marsupial cheekteeth. Ihni 18, 137-464. 

— Si W\ni , M. (1976) Results of the Ray B, Lcmley 
Expeditions, Pan I. The AHinglunu Inrmation and a 
new Pliocctlc vertebrate fauna from northern 
Queensland, ihid 17, 179-397. 

Bahmtii, M. .V ftl 1 1 n. H, [till) Prehistoric man ui 



Lake Mungo. Australia, by 32,000 years BP, Nature 240, 

Bartholomai, A. & Woods, J. T. (1976) Notes on the 

vertebrate fauna of the Chinchilla Sand. Appendix 2 

in Bur. Min. Resour. Bull. 166, 151-152. 
De Vis, C. W. (1887) On an extinct Mammal of genus 

apparently new. Proc. R. Soc. Qld. 4, 99-106, pi. 1-4. 
(1888) Notes on the genera Zygomaturus and 

Nototherium. Ibid 5, 111-116, plate. 
____ (1891) In confirmation of the penu* Owenia so- 

called. Proc. Linn, Soc, \>.SM <Ser 2)6, 159-165, ; i 


(1895) A review of the to-il - of tne 

macropodidae in the f '■■-■■ rul; niMaj i>m ibid 10, 

75-134, pis. 14 18. 
Stirton, R. A., Tedford, R. H. & Woodburne, M. O. 

(1968) Australian Tertiary deposits containing terrestrial 

mammals. Univ. Calif. Pubis Geol. Set. 77, 30 pp. 
Tedford, R. H. (1967) The fossil Macropodidae from 

Lake Menindee, New South Wales. Univ. Calif. Pubis 

Geol. Sci. 64, 156 pp, 4 pis 
, Williams, D. & Wells, R. (1986) Late 

Cainozoic sediments and fossil vertebrates pp 42-72. 

In Wells, R. T. & Callcn, K. A. (Eds) 'The Lake Eyre 

Basin — Cainozoic sediments, fossil vertebrates and 

plants, landforms, silcretes and climatic implications". 

Australian Sedimentologists Group Field Guide Series 

Hi 4. (Geo!. Sex Aum.. Sydney! 
tt<.oj\v .1 T (I960' frifctiliferauti ffoviatilc and cave 

dope cts. In Hill. U «\ Derimead A. K (Edsj ! he 

Gool :v ol Queensland G ai S<n. . !<«/. 7, 393*403. 
(1968) I he UlemiLj ^i the &viinct marsupial genus 

Nototherium Owen. Mem. Qld. Mus. 15, 111-116, pi. 





VOL. 113, PART 4 



by A, Z). Austin & G. R. Allen* 


Information is presented on the large complex of hymenopteran and dipteran parasitoids associated 
with Uraba lugens Walker (the gumleaf skeletonizer) in South Australia. A key to the 22 species 
involved is presented, along with notes on identification and relationships with their host. Two 
species of microgastrine braconids are described, Cotesia urabae sp. nov. and Dolichogenidea 
eucalypti sp. nov.; both are parasitoids of the larval stages of U. lugens. 

KEY WORDS: Uraba lugens, Noctuidae, parasitoids, hyperparasitoids, Braconidae, 
Ichneumonidae, Aphelinidae, Chalcididae, Elasmidae, Eulophidae, Eurytomidae, Eupelmidae, 
Trichogrammatidae, Tachinidae. 

Transactions of the Royal Society of S, Amsu <W9), 113(4) W 184. 




by a. D. Austin & G. R. alj_en + 


Austin, A. D. St Allen, G. R. (I9S9) Karadtaids of Uraba tu%ens Walker U.epidoptera: Noctuidae), wiiti 
description of rwo new species of Bracomdaa Trow, H Sov.S, Aust. 113(4). 169 IS4, 30 November. 19S9. 
Information is presented on ihe large complex of hymenopleran and dipteran parasitoids associated wilti 
Uraba lupins Walker (ihe gum leaf skeletoni/er) in South Australia. A key to the 22 species involved t< 
presented, along with notes on idcni»ficaiion«H;d relationships wlih iheir host, Two species of niian^aMriue 
btaconids are described, Cote^ia untbue sp. tfCK *nd OuUchogen idea eucalypti >p. nov.; bmh are parasHoids 
of ihe tai val stages of U. tugeas. 

K^y Words- tMth(i tuft/ens t NociuWfts, paravtoids, tryp<rpaiasitotdi, Biavomdae, Ichneumonldae, 
Aphdinidae, Chalcididae, Ela.\midae, Fulophidae, Kuryiomidae, Euueimidae, Trichosramnwlidac, Tadunidac. 


Urate lugens Walker, the gumlcaf skeletonize*; 
is a native noctuid moth and has been collected 
from all states ol Australia (Turner 1944), It has 
been recorded as damaging stands of eucalypt 
.species in eastern Australia (Brimblecombe W62; 
Campbell 1962; Harris |972\ 1974; Harris ef t?/. 
1977% the Adelaide region and south-western 
Western Australia (pers, comru. F. D. Morgan; 
Strelein 1988). Occasionally outbreaks of this 
species can defoliate large areas "of native forests. 
Several such our breaks have occurred in stands of 
liucutpytus comatdulensts Oehnli. along the Murray 
Valley region of N.S.W. and Victoria, where, on at 
least lour occasions, morelhan J0,(KH) ha have been 
affected (e.g. Campbell 1962; Harris 1974; Harris 
ei nl. 1977'). Apart from the widespread damage 
a.vsnciated with this species, it Is also responsible 
for the partial defoliation of individual eucalypt 
tree* planted as ornamentals In parks and gardens. 

Other than ihe work ol the above authors and 
those of Morgan & Ccbbinah (1977) and Cobbinah 
(1983), very little has been published on the biology 
and ecology of U- lugens, while even less ha*, been 
reported on its parasitoids. Brimblecombe U962> 
reported five species of primary parasttoids as 

! Harris, J. A. 0972) I he effect of flooding on population 
density of die gam leaf skeletoned' morh, Uraba iuge.'v, 
Wulk-. in tiarmah Stale Hires! . Knvsl Commis-si«ii. 
Victoria. Research Branch Report. No. 25 (unpubll 

•HaTris, J. A. Neumann, F. G. & Ward, B. (1977) An 
onlbteak of the gum leaf 4:e!cronizcr. Uraha ittgetts 
Walker, in river red gum forest near Barman- K>reM 
Commission, Vicloria. Kcscarch Branch Report, No. 
87 mnpnblj. 

Department of Hnromolopv. Wane Agricultural 
Rematch insnmte, Univcrslry of Adelaide, Glen 
Osmond, S. Aust. 50M 

attacking U tugens and Campbell (1962) ten species. 
However, in most cases these parasitoids were no! 
identified I ml her than family level and, if so, their 
identification was not reliable. One of us (G.R.A.) 
has recently completed a major siudy on th<"- 
interaetion of this insect and its parasitoids in tin- 
Adelaide region. This work shows thai the 
immature stages of U. lugeris support a diverse 
complex of hymenoptet an and dipteran parasitoids 
(22 species — Table 1), which includes both primary 
parasitoids and hyperpatashoid.s. In this paper we 
provide a laxonorn're framework for the information 
on the behaviour and ecology of this parasitoid 
complex and its interaction with U fUge/Wj which 
will be published elsewhere by G.R.A. Here we 
present a key to identify alt the parasitoid species 
involved, and provide notes on their taxonomic 
posilion and biology, including information on the 
stage attacked and place of pupation. Two of the 
mote common species reared from U, lugens larvae. 
which are members of die braconid subfamily 
Microgastrinae, and are the subject oi detailed 
behavioural studies by O.R.A., are described hetv 
as new. 

Materials and Me* hods 

All life history Mages of ti tugens were collected 
from euealypts (mostly E, canialdutemiy. C 
kwoxylon F Mucll. and £. microearpa (Maiden) 
Maiden) between 1V85 and 19S8 at several sites 
within 7 km of Adelaide G.P.O, They were brought 
to the laboiatory, held at 20°C in rearing cages, and 
larvae provided with new foliage, so that any 
parasitoids present could develop and emerge 
normally. Parasitoid cocoons found with their dead 
host in Lhe field were also collected and reared in 
Ihe laboratory at 20°C until adults emerged. 

I/O V !> AHS11N K it K. At 1 EN 

I A. 141 h I. Summary of /vhttionshipi between U. lu.ecns and Us parasitokh ana' hypcrparusitOHh 



' (P) n- Hypei parflGHflid (H) 

trl /wetvn 

-;u^c Stage tfftt!jj$ 


Family Sotiiars (S) 01 (jf^an-nis (( ii 



rnrfntyjutntna sp, 





CoU'Utt tfrflft« 





Di linlw/tr'Mh'a fiwufyptt 



1m va 


fr'uftiwirwi sp. 





CijsWiiila intern 



liit\ J 


Lwtistif ,ftff\tcejte 



(ill vfl 


Crthorw; '.p. 





Xaritfu >i>ttt}pt<i rh opulnceim 





A'tlrocephaht* *p 





ftrurhvm'rut ip, I 





Wmihcmta laren/IU 





Lur}ivtfiu ^p. 




pupa /para-moid cocoon 

Centrodom vy 




pMrMMUnd cocnon 

U>ut:h*mvrHi sp. 2 




paranoid cocoon 

hlrtstuus au^ruhemu 




pamitoid cocoon 

mwirt imkt 




parasitoid cocoon 

t\vitohus lip. 




parasitoid cocoon 

species indei. 



parasiuiid cocoon 

AnpMunis sp_ 




pma^iioid cocoon 

\U\wchoru* $p r 




paia-idiifl c^ooon 

Pawphvlax Sp. 




parasite-id cocoon 

pfemtiujfitsi c .p- 




patasitmd cocoon 

Parasitoidi were stored in a freezer or in 70^0 
cthanol prior to mounting on pins or card points, 
Material for S.fcLM study was washed in hall" 
strength concentrated liquid soap, rinsed in distilled 
water, dehydrated in an alcohol series and critical- 
point dried using an Fmseope CPU 750, before 
being examined under a Cambridge Stereoscan 250 
(Mk Mi) electron microscope. Terminology for 
morphology follows Boueek (1988), Fady 
(I96X), Gauld (1984). Harris (1979), Mason (1986) 
and van Achterberg (1979). The term l alitrunk ! is 
used for the tfeorOK pl«5 propodeum, and faster' 
is used for the post-propodcal segments, The 
abbreviation *T' refers to the gastral tcrgiles. 
AbbrevialiOus for collections are; ANIC\ Australian 
National Inseci Collection, CS1RO, Canberra; 
WAR1, Waitc Agricultural Research Institute, 
Adelaide. Voucher specimens of all species are 
lodged 111 the Waitc Institute collection. 

Ke> l« Hit* |)iirasiW»ids ttf t/fsbB tttggty 

in Sonln Auslruliu 

l\vt> |t|ijr ol' wings developed; clonal surface very 
rarely with -lout burlier uusp-likehi appearance [ovi 
posiror always developed in female and nsually clearly 

visible (Figs 5, 7, 291) (Hymenopicru) 2 

Only one pjiir-tif wine 1 - developed (foa 1 WlOga}; dorsal 
surface with numerous sioui. brinies; blowfly like in 
appearance (Fij4> -18. 49) iscukim with several block 
loncinidinu] band*) (Tachinidae).. , , ...2! 

foa wtog wiib reht ivory complete venation (e.g. Fig> 

I, :, h, 19), 3 

Fore wins wilb pij:rneniedvenaiion reduced loanicnur 
niurytii (e.e f'l^s 21, 23, 27, 39) -.-9 

I, Foa- winy, h trh venation disiat io piejvjsnf'.ma wanting 

(Fks t, 2)\ venl 2(11011 nlweni (Kraconklaej 4 

Foic win?' uidi dLsi.'it \ein> presenl and welt 
piyincnied; \ot\ Stracu prc>eiu (H^i 4. 20) (Ichneu 
momdaet . .$ 

4. Fropodeum with loneiiudinal medial carina, coarsely 
seutptmed .*u fcqst anteriorly (Tig. 12); legs ted lo ted 

yellow ( 'oti'sict uruhae sp. no*. 

Propodeum wirb large earmale ^recila and horiAMital 
carinae extending to lateral margins ot propodeum 
(Fig. 13): legs dark brown |o black with distal parts 
icddish Douchngenirff'a eucalypti sp. nov, 

5 Fore wing with an areolet (Figs 4. 6). , , , A 

Fore wing without an areolet (Figs 19. 20) 7 

6, Senium md prnpodemn coarsely punctate or 
ru^m'osi.: ovi|io^itor vciy -linir fl-ttv S); rt yemtwlKi 
wilhom tontirods proirudin.y posteiiorly (body dark 

biown io black, le^s icddish), , , . 

. Ctishttinti wit-nt lei man & (iauht 
Scutum and pmuodeum t J, -*neiaHy un^eiilpuned 
Uwcepr lor propoJeal carinae); oviposiior cMcndiirc 
well p..;is| ptHienor ^nstef (lie. 1\\ >* feniraha with 
pail ol Vjan iods(ront;v>qnama)pn,>I'it(liMu puvtfiit.nK 
(hodv yellow-brown wirii Jaikci maikiiij.'.si, , . . . . 
frfvsochtiriis sp 

T, Body bn^hl yellow with hl/kk niiidtiiL's: 1 1 Mion timl 

bttfild basaliy (Ji^. 16) 

Xanthnptmphi rhopt/incctw Krieeer 
Bodv not so colomed; Tl uantm l.u^ally [Hfc> 17, 18) 


8. Fore winu wilh radial cell sborl and broad (Fi^. 19): 
I \ tint, broadening distally (Fig. 17) (small species. 
length 2 1—1.2 mm not including § ovipositor; 
body black except tor T2 which is ye) low-brow nj 
Paraphylax sp- 
here wing wuh radial cell lone and narrow (Ftp. H)). 
mhnlai in basal hall, bulbous in tlisml hall' (F'ip 
IK) (large species, length 6.5-S.7 mm not inclndinc" 
ovipovdor tor v: head and aliirunk black, easier 
reddisli-luown . , . f-riftonn sp 

V. lemur 01 hind leg greatly expanded, luulned 
01 senaied Moug lower margin (Fiji. 24) «..M 

1 ididtie) _, , , jo 

Femur ol' hind leg noi mal, smooth alon& lower margin 
(FigS 2S, JS> /_ (2 

10. loiv wing wilh marginal Will ffliwta longer 
hOMmarjuiial vein (Ha. 22); upe\ oi' hind tibia tapmim 

into sirong spine (Hrachytttc/M) , , , (I 

I ore wing with marginal vein about s«mc length as 
posimaiginal vein (Fig. 21); apex of hind tihi>* 
perpendicularly nuneaie (Fig. 24) (huge species, 4.8 
mm in length; body black, hind leg dark red-brown 
marked with black I .... Antrowphatut $p, 

1! Kody black with rtd hind lemur and iihu;4 2-4 4 mm 

jj ienyi h . 8wchv*, n -nu sp I 

Hodv I.IiilIv with vvlutc-yeliow mmkine-s on wivtlfeu :m<J 
k *L'v ($r$3 mm in Imgth Dradn/thYi* >p. 2 

pah asiioids or ukaha rxtCSMS 



Hind v to\a developed os huge flat disc; hind ohm w.|h 
» limine! erkveross pattern of setae (Hg. 23) f lot r wing 
wiih itjpial vt-'tn very short (tie. 23). body dork. 
tcaitl.i ninl leys except for hind cm*a pale; bodv length 
ot v J. 8-2.7 mm, of 1.3-1.9 mm) tntovmidae) 

- • - Eiuwtus ausjiatn-nns i.iiraull 

Hind Cflwfl, hind libia and siigmal vt-in not as above 


F'ronoium (seen doisuUy) large and quiUUaiigutar, 
dorsal surface ol* alitrunk coatvetv sculptuted (tig. 
2A), foie wing venation oi in Fie. 2~ (body bluet; and 
non-metallic, legs with .some pale 'markings) 

lliuryioimdiie) Eurytoma kr. 

Pronotuui not large and quiidmntmlar and alitrunk 
without such sculpiuim^: body'Wien jlKtftlNC in 
colour , , 14 

M, Hody length greater than about 1.5 mm; >»>isier 
M-|Mruied from alimmk by narrow tfaSfcl (fogs 28, 29* 

, 15 

Uody length [vtti than !.<"• mm (minute specie^; paster 

hrotidly attached to iiltrrunk oi appearing so (l\n. 4*1) 

-. .... 20 

15, Tarvi 5-segmemed <ef J'ig< 24, 25) .if, 

Hirsi 4-sL-gnienicd (Tig. 35) . 18 

Ifi Mesopicurnn nor enlarged and shield-like (Kg, 2tf); 
DQdv rUlflej rohusi with largt: head and alitrunk ( g 
easier iii lateral view -dimply angled: cr jt&xt&t xmall 
arid flattened; body length of '-'■ 2.3-3.3 nun, 

1.7-j 4 nin , tritiaiiK-, ^^ i:u l(1 co [ our Ult |j 

yellow-hmuvu leg* and antennae) (Plerom/duhie) 

Mcsopleuron befit mid shield like: bodv sumewhat 
elongate <Hg. 20) (Fupclmidae). . . .. .' \i 

17. Dorsal snrlaee oi sculum flnrrenc/d wilh railed uiuvr ior 
timinmlar area (big, Iljj fore wing with broad 
pigmented bands the. 30) (mosllv dark m ootout vtflfl 
tnetallie green unite; body 2.1--2.5 mm in length) 

• ••'■'- * --.»,,i Anastutus \p 

Dorsal surface of scuium not particularly tlatteitcd, 
without anienur raised area <l-tg, 32); lore win^s. 
hv.ilme (head and alitrunk hi ight metallic green, ga.;tet 
darker). . . . . ,gen. & -species indei [qp only) 

18. Fo r e wing w)ih stigmal and posimniiunal veins very 
shon (I iy. 34); ;interioi' s^ulcllum longttudiuallv 
striate (Fig. J3) (hody daik wilh mcmllk y'reen tin^e, 
Fl-i.5 mm m h-ngi.h), ... Miolnus <,p 

1 ore vvntg with u tonal und posiinni«inaJ veins lon t - 
(Fit P); ^culellwm Jtnooih or with fine lenuiksic 



Dorsal head and *hiinnk with scattered ti3n% btte&c 
like hmrs (fig- ^n) (body movtly dark and non- 
metallic; ryes red; antennae, lee^ and broad patch on 

dor.'Ol Ba.ster vellow-hrowii). , , FMf>!ecim.s sp. 

Dorsal head and alitiunk with :;hon<:i ["im;r hain 
(dorsal aliiruuk with distinctive rm-tulljc yreen and 
vellow niiii kings tFig. \X)) .& n . &. species mdei 

2(1. Tarsi 3-u-gmented ttiy 40); antennae S-segm^nted 
(hip 41, 42); tore wing veTy hrinitl (Fig. JO) 

iTrichooiammatidae} . • , Irichtwonmhi sp, 

Tiirsi 5-scgmeuted (Fie.. 47); antenna^ 6-segmcmed 
(PigS 45. 46); foiv wing narrowct (fjg, 43) (Aphrl 
initiac) . , , , , CkiHtrndmu sj.. 

-I. Hairs covering ocopm (posfctior part of heatl) silver 
gtey; abdomen in dorsal view wuti mitcuoi; 
and posterior parts black, lateral areas hmwn (Fi;;. 

•**)- ■••■«.' Wiftthama taterults (Mat-quart) 

Hairs coveiine oeciput yolden brown, 1st segment oi 
abdomen black, other segments bla.;k with parches 
of Mlvei (Fie. 49). . .h.Korhtu ftuvnvps MlMSifUHH 

frcatnieni ftf species 

loittily Bracotiitbic 

FWh 2, X, 10-tl 14 

Uahtype: 9, ANIC, S. .\ust-, Adelaide (Mitchmn). 
tcaied c.\ Uraha htgetm un huajtypws miaveurpa, 
l-oIF I4a.J985, erncie h ed 3.\il9iSs. {}, R. Allen 
Paratypes; S'? o, 2(FT.-y, same data as holoiype 
cscept some with different suburbs of Adelaide and 
different dates; 179 ffj 13 er rr, S. Aum. ( Adelaide 
<Waite lu.Miiule campus), various coHeciin^ and 
emergence dates duting 1964, I . Hope. 5 v $'<}&> 
S. Anst , National Park, Bclair, cull. ZO.xi.1%4. 
emerged 25.xi.l964, F.D.M. (5^ ?, $&«r, ANIC; 
rest of material including l o go J coated lor SCM 
(wings slide-inoun;cdr in WAR ). 


length. 2.9 mm (2.5-3.2 mm. t -= 10) including 

Colour, Body black; palps yellow; legs except lor 
co\a yellow-brown, hind leg tnluscuie Jisially, dislal 
cud ol hind tibia with dark pylch; anterior pleural- 
s:urnal region of paster dark red. \\tnys hyaline, 
siiema cvenlv and darkly piyjiicnlcd as is rcsi ol 
foic ftlllg venation. 

Head, In anterior view vertex arched so thai hea<i 
is somewhai circular; face and gain with loiiiiHi 
^hite hairs and associated I'irie colliculaie sculp- 
turing; inner margins of eve* Virtually parallel; in 
dorsaJ view ocelli in wide itinriglc. posterior tangem 
ol median ocellus coincident with imaginary line 
across (he anterior margins ol lateral ocelli; lions 
and medial occtpui smooth and hairless; temples 
wnlt white hairs and associated oatlKularc sculp 




tlgfl 1-7, 1-2, hire and hind wings; I, DoHcho^enuwa eucalypti ^p. npv„ 9; 2, Coieva urabae sp, nov.. 9 ; 3, Lateral 
view ol' gasier 01' DatiL-fio^eructea eucalypti sp.nov., 9; 4, Distal tore wine of \fe,%ochon<s sp., 9; !H5i Cusirwna 
micro ■ Jcrrnan & Gauld, O; 5, Lateral view of \gasicr; 6, Rire wing; 7, Lateral view ol easier ot Mesochurus 3 p r , 9 , 
Scaler Htu> I, 2, 4 ami ft = 0.5 mm: Fig. 3 = 200 ft©; Figs 5 and 7 = 1.5 mm, Abbreviations: a = areolet; pt = 

Hiring which N slightly coarser than on face: vertex 
with few scattered short hairs otherwise smooth; 
antennae slightly longer than body, distal 4-5 
segments only slightly longer than wide. 
Alitrunk. Scutum punctate with covering of shortish 
hairs, punctation denser alone courses of notauli 

and along lateral margins (Mg. 8), punctation along 
posterior margin becoming slightly longitudinally 
elongate; courses of notauli faintly depressed, these 
faint depressions broadening posteriorly; scutdlum 
faintly punctate; phragma of scutellum exposed 
along posterolateral margins (Fig. 8); propodeum 



Figs B— 11, 8-9, Dorsal view of alitrunk; «, Cofesia urabaesp. nov., 9; 9, Dotichogen idea eucalypti sp, run <?• 10-11 
Coiesia urabae sp. nov., Q; 10, Dorsal view of gaster; 11, Lateral view of gaster. Scales: = 200 «fn, 

i :.; 

\ |>. AUSTIN &G, R M I IN 

coarsely rugose to rugose-slnate in anterior half, 
generally smooth wilh faint rugose punctatitm in 
posterior half; medial longitudinal carina welJ 
developed with associated short horizontal and 
oblique eartnae (Fig, 12); mesopleuron finely 
punctate in anterior half with associated short hairs, 
smooth and hairless in posterior hall' except For 
compact group of 5-6 foveae medially; metapleuron 
smooth anteriorly, rugose-punctate posteriorly (Fig. 
14); hind coxa faintly punctate on dorsal surface 
with associated short hairs, this sculpturing 
becoming coarser on ventral surface. 
ttmgs. Fore wing with veins r and 2-SR sharply 
angled, r slightly longer than 2-SR, cu-a almost 
striate; distal pan of basal cell and anterior pari 
oT sub-basal cell devoid of hairs or almost so; discal 
cell sparsely covered with hairs; hind wing with vein 
r (spectra!) present (Fig. 2). 
Caster, TJ as wide as long, broadening posteriorly, 
surface in posterior two-thirds coarsely puncialc ro 
rugose-punctate, becoming coarsely striate in 
posterior one-quarter tFig. 10); scleratized part of 
T2 rectangular, slightly wider than Tl, coarsely 
rugose-punctate with a few longitudinal stnations, 
sculpturing fading to nearly smooth in postero- 
lateral corners, longitudinal midline smooth; T3 
usually rugose-punctate in anterior one^quarter to 
rwo-thinis, with scattered hairs, posterior part 
smooth, in some specimens T3 virtually smooth 
throughout but always with a( least anterior margin 
with band ol punctaliom rest of termites smooth 
wilh scattered hairs, in lateral view hypopygium 
pointed, extending past posterior gaster, with scat- 
Li:icd fine hairs, ventroapical margin not indented 
(Tig, II); ovipositor sheaths with few apical hairs. 


As for female eseept for length, 2.R mm {2(y-} i 
nun, n = 10) and sexual differences ^genitalia and 
development of hypopygium). 

Other material examined EL Aust., suburbs of 
Adelaide, various dates and collectors, 100 9, 
Ifrd cr -WAR] (excluded from type scries because 
material is damaged or inadequately labelled). 

Comments: The scuiplunng on the propodeum and 
TI-T3, the shape of these sclerites. the form of the 
hypopygium and ovipositor, clearly place this 
species in Cotesiu Cameron. Cotesia has previously 
been referred to as the glomerulus species-group of 
Apanteles s.L (see Mason 1981; Nixon 1965), and 
is the latgesl generic level taxon in the subfamily 
Microgastriuae, ibc lattet comprising some 1300- 
1500 described species world-wide (Mason 1981). 
]|] Australia Cotesia is common and diverse, but 
other than several species introduced Tram Europe 

and North America as bio-control agents for certain 
lepidopieran pests (viz. C Jlavipes Cameron, C 
gtomerutu (L.J C. ka7.ak ( Telenga), C margin iventris 
(Cresson), C plutellae (Kurdjumov), C rutwula 
(Marshall) and C. tu/urus (Hahdayj), the 
Australian fauna remains unstudied. A few poorly 
characterized species, which presently remain under 
the name Apanteles $,L may turn out to belong in 
Cotesia, These species are unlikely 10 be conspecific 
with the present species as their type localities are 
outside the known distribution of C uraboe sp. nov. 
or they are associated wKh other hosts. There arc 
no workable keys to Judo-Australian species of 
Cotesia. However, the key In Nixon (1974) to the 
north-western European fauna can be used to 
separate C. urabae sp. nov. from four of the seven 
introduced species mentioned above Of the other 
three species, G Jlavipes, is very different to G 
untbaesry, nov in that its body is strongly flattened 
dorsoventnilly (see Austin 1989), while C. kuzuk and 
C marginiventris can only be identified reliably in 
association with their hosts, Helttoverpa spp. and 
Mythmina conveeta (Walker), respectively. 

Biology; C, urabae sp, nov. is a solitary, primary 
endoparasitoid and oviposits into early to inter- 
mediate larval instars of its host, emerging from 
intermediate to late insrars before pupating. The 
pupal cocoon is alongside the host and is sulphur 
yellow-green with a surrounding silk matrix. This 
species lias only been reared from U. lumens. 

DoUehogenfden eucalypti sp. nov. 
FIGS I, 3, 9, J3. 15 

Holoiype: 9, ANIC, S. Aust-, Adelaide (Wighgate). 
reared ex Uraba lugens on Eucalyptus camal- 
dulensis, collected l.\i.l985, emerged 17 xi.1985. 
G, R, Allen. Paratypes: 15 9 9 . 20 j* a* same data 
as holotype except some with dilfercm suburbs of 
Adelaide and different dates; 13 9 v, 5c o\ S, 
Aust-, Adelaide (Waite Institute campus), \atious 
collecting and emergence dates during 1964. 1.. 
Hope (59 V. 5crcr, ANIC; rest of material 
including I 9 gold coaled for SF.M (wings slide- 
mounted) in WARt). 


Length. 2.9 mm (2.5-3.1 mm, n - 10) including 

Colour, Body black, wings hyaline with darkly 
pigmented venation, palps brown, (eg*, black with 
distal fore legs, tarsi of mid and hind legs and tibio- 
femoral joint yellow-brown. 
Hftid. Mostly smooth except for line collicularc 
sculpturing associated with dense covei Iflfi of short 



Figs 12-15. 12-13, Propodeum and first and second tergites of gaster: 12, Cotesia urabae sp. nov., 9; 13, Dolichogemdea 
eucalypti sp. nov., 9 ; 14-15, Lateral view of alitrunk: 14, Cotesia urabae sp. nov., 9 ; 15, Dolichogenidea eucalypti 
sp. nov., 9. Scales: = 200 /*m. 

1 1* 


hairs; surface *iili characteristic dull lustre; in 
ameflor view inner margins nt eyes virtually 
parallel; in dorsal view ocelli forming wide triangle, 
posterior tangent of median ocellus coincident with 
imaginary une across the anterior margins of lateral 
neelli; fine collktilatc sculpturing and associaled 
piJosity slightly denser across vertex and occiput, 
except for smooth narrow band around posterior 
margins of eyes: antennae reaching to posterior 
gasicr or slightly beyond, drstaf three segments 
slightly longer than wide and sometimes slightly 

Atitrunk. Scutum coarsely punctate with punctures 
mostly closer to each other than their own diameter, 
except along posterior border and along courses of 
notauU, which are thus faintly indicated (Fig, 9); 
scuiellum smooth; scutum and sculellum densely 
covered wnli Short hairs; melanolum rather broad, 
antuolateral margins emarginaie so iiiai phragma 
of scutellum welt exposed; carinae forming 
prnpodeal areola raised well above surface, eamiae 
extending laterally below horUomal midrine> these 
carinae with dorsal and ventral extensions forming 
aistuiac, but not enclosing spiracles (Fig. 13); 
anterior part of propedeutic mostly smooth and 
setose, posterior part wiih faint rugose-punctate 
sculpturing which becomes more obvious laterally; 
mesopkim.n fttOSC in Ulterior half, smooth 
posteriorly; metapteuron mostly smooth, except tor 
venuopoNiuioi one third which is rugose-puticiare 
(Fig. 15); distal ton: tarsus without spine opposed 
to larsHl claw 

Wtttgsc Hue wtog venation -a? in Fig. I; costal and 
basal cells b*re posteriorly, hind wing broad; vein 
t-SC+R uecply bowed; r present bui faint; 
eubilellau cell moderately broad, suhmediellan cell 
rminded posieriorlv. 

turner. "II as wide as long, slightly widened In 
posterior Ml(\ lateral margins slightly emarginaLe, 
Mirfuv mostly punctate, .^nale-punclate along 
lateral margins and striate in posterolateral corners 
and along, posierior margin (frig. 13); sclcrotued 
ptrl of T 2 slightly wider ibanTI, 25 x wider than 
Ions mostlv smooth Willi I'ami scattered punctures; 
13 slightly longer frum T2 (14:11); T4r-T6 shorter 
than 12 (Rll); I ) w-iy Short, about onc-quancr 
leng* 6| 12; l *-T* all smooth; T2-T7 each with 
single transverse row of hairs; ovipositor and 
viH.iir.s shorl, not exiending far past posterior 
gastcr; ovipositor with strong distal attenuation 
1 1 Ig. M; hypoprygium lacking obvious lateral creases 
rtiflUgfi weakened normally in ventral midline. 

menicd in medial area so that it is transparent, flft$C 

of wing venation generally with less- pigmentation 
than female; H sometimes with dense rugose- 
punctate sculpturing merging with posterior 
striations, otherwise same as female except for male 
genitalia and Lacking bypopygujm, 

Comments: The sculpturing of Ihe scutum and 
propodeum, shape of Tl and T2, form of the 
hypopygium, and shape and fringe of the van rial 
lobe of the hind wing clearly place this species in 
Doltchogenideu Vierecfc- Previously Do/ichogemdea 
was considered as three related species-groups in the 
genus Apanteles s.l, t \U. the uttor, laevigatas and 
lon%i[)Qipis species-groups (see Mason 1981; Nixon 
1965, I967| This species falls into the ulior group 
which was revised by Nixon (l%7) for the Iptla 
Australian region. In this work D. eucalypti so. nov. 
keys out to D. cieo (Nixon) (eouplel 83}, a species 
known only from India and associated with a 
nyrnphalid host Erihoae atja Fclder, or with some 
difficulty it keys as D. caniae (Wilkinson) {couplet 
31). which is known only from Java arid associated 
with a limaeodid, Cantu bandura Moore I.Austin 
1987). Apart from having different hosts, these 
species differ from D. eucalypti sp. nov. in that D, 
caniae has an unusual striate sculpturing pattern 
on Tl and T2, and D. cfeo has the sub-basal cell 
evenly and darkly setose, the hind femur yellow in 
colour, the proximal half of the ovipositor very 
broad, and the mesopleuron coarsely rugose- 
punctate anteriotiy. This is the fourth species of 
Daitcho^enidea recorded from Australia, the others 
being Q lipsis(Si\on) comh. nov.. D, miris (Nixon) 
comb. nov. and D. tasmanica (Cameron) comb. nov. 
fcxaminaliori of the holotypcs of these gpiacfefl shows 
that they differ from (X eucalypti sp. nov, in a 
number of important characters. All three have the 
ovipositor much longer, being at least as long .is 
the hind tibia, and, in addition, O, tfpsis and D. 
tasmanica have a white spot on the cheek and a 
much reduced propodeal areola. These specie- car 
be readily separated from & eucalypti sp, nov, using 
the key in Nixon (1967). 

Biology: D. eucalypti sp. nov. is a solitary, primary 
endoparasitoid Avhich oviposits into early to 
intermedial? larval instars of its host, emerging from 
intermediate to late insiars before pupating, The 
pupal cocoon is spun alongside Ihc host and is white 
in colour and lacks a surrounding silk matrix. This 
species has only been reared from K lugens. 


.V. Km female csccpt as follows Length 2.7 mm 
(2.5 LB mm, n x 10); aliuunk very sligh'ly 
flattened doistm-ii. tally; fore win a stigma Unpfe- 

Family khneumuiudae 
Xanthoptmpla rhopalocems Krieyer 

no. 16 


This species Is eavily identified by fts distinctive 
colour pattern and fore wing venation, in South 
Australia il is known from two specimens {WAR! k 
one collected ;n an adult in the Adelaide region, 
the other reared from U. lugens at Keiih. It is also 
known from Oueensland and 'Tasmania where it has 
been recorded from U. lugens, and the tortricids 
Epiphyas posivtttanu (Walker) and Merophvas 
rfivutsam (Walker) (Brimbleeombe l%2; Dmnble- 
con 1940; Gauid 1984; also see Townes & Chui 
1970J. It is a solitary, primary endopatasitoid and 
emerges from ihc* pupal stage of its host. Sec Gauld 
(]yK4) fur additional laxonoroic information and 
list of synonyms. 

I*araphvlax sp. 
PIGS'r?, 19 

Paruphytax is a large Old World genus with more 
than 50 recognized species from Australia, (he 
majority of which are unde.seribed (Gauid 1984). 
Where their biology in known Parnphyktx *pp. have 
been recorded as primary and hy|>eiparnsitoids\ 
mostly of lepidopteran hosts, Apart from the 
characters given in the key this species is notable 
in comparison to other parasitoids associated wilii 
U tugem for its relatively smooth unsculplured 
body (except for propodeal carinac) and lateral tecih 
on i he propodeurn. The species here belongs to the 
corvax species-group (sec Gauid 1984) and is only 
known from the Adelaide region, where il has been 
reared as ui obligate, solitary hypcrparasnoid 
through C urahoc and P. eucalypti 

Erihorus sp* 
FIGS 18, 20 

Ihis is a distinctive species when compared to 
ihc other ichneumonids associated with U, fuvefis. 
In addition io the characters given in the key ibis, 
species has distinctive reticulate-punctate sculp- 
turing on the scutum, seutellum and propodeurn. 
Erihorus sp. is a solitary primary pnrasitoid pjj V. 
lu&ens. ovipositing into the larval Stages and 
emerging from the pupa. Jt is only knrmu from the 
Adelaide region and has been reared from its host 
On various occasions since J%5 (WAR1. unpub- 
lished records). 

Casinaria micro Jerinan & Gauid 
FIGS 5, 6 

This species is a solitary, primary endoparusiioid 
easily recognized by its lore wing venation, shun 
ovipositoi and colour, It has been recorded I torn 
all stales in Australia and, although it has been most 
commonly associated with U w$a*S, C- micru has 
been teared Irom species belonging ro three other 

distantly related lepidopteran families — Geomct 
ridae. Oceopltoridae, NotodoiHidae (see Gauid 
WK4; ierrnan & Gauid 1988). The pupal cocoon is 
constructed ncai io (Jerman & Gauid 1988) Bi 
underneath (observations in this study) Its dead 
bl val host and is attached firmly to the leaf sut lace. 
Tt i^ grey-brow u and marked with characteristic 
black spots. See .Ierman & Gauid (1VK8) lor add- 
itional taAonomjc information and list of synonyms. 

Mesochoru.s sp. 
FIGS 4, 7 

This is a large cosmopolitan genus of primary 
and hyperparasitoids oi lepidopteran and coleop 
teran hosts; most Indo-Austtaltan species arc nndes- 
enbed (Gauid 1*84) The Npecies recorded here is 
a solitary obligate hypcrpurasitoid of U. lugeni 
through C, mtcru, C umhoe and D. (fiav/ypti, and is 
known only from the Adelaide region. It is a pale 
coloured delicate species with fine thread-like 
antennae. The male is distinctive in having the 
gonoso,uama oT the genitalia extending from the 
posterior gaster as a pair of (oils tods, 

lamilv Cludcidiihic 

HnnhvmerUt spp. 
no, 22 
This U a large genus in Australia wtrh nearly 70 
described specie^ the majority of which ale 
primary, pupal parasitoids of Lepidoptera (Boucek 
19-S8). The two species recorded here (both 
unidentified) can be separated easily by the 
character m the key. Species J is a solitary, primary 
parasitoid of U. itt$M and emerges from the Iiom 
pupa. Species 2 is solitary and hyperparasitic 
through ( . urabae and iX pucutypti t though it b 
not known whether this relationship is facultative- 
or obligatory. In eastern Australia two species of 
liruchymerk have been reared from V. lumens, w/. 
tLjffQ&gtiUl (Cameron) (Rmnblceombe l%2) an<l 
Ji ruhripes Ginuiit (Campbell 1962) [B. rubn'pts is 
considered a junior synonym of H, Wuta (Walker). 
see Bouc'ck 1988). However, the material from these 
records would have been identified ai a time when 
the Australian species in Ihc* genus were contused 
by most authors and hence might be misidentified, 
These names .should thus be used with some care, 
especially since InmcVk <|9XX> did not sex.- unv 
matciial in Australian collections or elsewhere 
reared from U lupens that he could assign to eithci 
R .froggtilH or B, teuta, 

Atitrocepiuilw) so 
I IGS 21 M 

Ibis species is- lep-e^ulcd here by a single 





specimen (eared as a primary patSSltofd from rhe 
pupa pf U. Itigens in (he Adelaide region, !| is easily 
distinguished by the characters in the ke.v. There arc 
more than 60 described Australian congeners which 
arc discussed by Boucek (19X8). 

Family Eurytomidae 

t'utytoma sp, 
DOS 26, 27 
I his is a large cosmopolitan genus whh more 
than 60 described Australian species (BouceK 1988). 
Biologically ihe group is very diverse including 
phytophagous species, primary parasitoids (mostly 
of lepidoptcran hosts) and hyperparasitoids. The 
species recorded here develops as either a primary 
grcgaiious parasitoid of U. lugens, or as a solitary 
hyperparasitoid through ( . urabae. 

Kamily Pleromulidue 

Ptemtuatus sp. 
Kit i. 28 

This species rs relatively easily separated from 
other t halcidoidea associated with Li lugerts by its 
robust body and metallic green colour. The genus 
is twxonomically very complex and the Australian 
species are in need of revision (Boucek 1988). The 
species recorded here is an obligate, solitary 
hyper pnrasiioid reared from Q /w#^v through Q 
micm C imbue and D, eucalypti in the Adelaide 

Family Klasmidae 

Elasmus austmliensis Girault 
FIGS 23, 25 

A distinctive species recorded here as an obligate, 
solitary hyperparasitoid of V lumens through C 
mtcru, C urabae and D. eucalypti. Previously it was 
known ro be hypcrparasilic and occasionally 
gregarious throueh an unknown ichneumonid 
associated with U. fugette in the ACT,, and also 
lias been collected from N.S.W. t and northern and 
southern Qld (lypc locality: Gordonvale, Qld). I'his 
t-s the (list record of £ amtraliensis from S, Aust., 
indicating that it is prohably distributed throughout 
south-eastern Australia. See Riek 11967) for 
additional la.xonomic information and list of 

Family Fulophid^c- 

Eupleclrus sp. 
FIGS J6, 37 

Euptectrush a cosmopolitan genus of gregarious 
ectoparasiloids of lepidopteran larvae, which is 
represented in Australia by 13 described species 
(Boucek 1988). The species recorded here is solitary 
and only known from the Adelaide region. It 
attacks the early to intermediate larval stapes, killing 
them before they pupate while pupating itselt 
underneath the dead host. 

Pediobius sp, 
FIGS 33-35 

This is 0) large, cosmopolitan genus of primary 
and hyperparasitoid species that attack a wide range 
of insect groups (Boucek 1988). There are more than 
30 described Australian species, most of which do 
not have associated host information (Boucek 
J 988). The unidentified species recorded here is 
easily distinguished from other Hymenoptcni 
associated with U. lugensby its distinctive venation 
and sculpturing on the scutellum. It i* represented 
by four specimens (W'ARI) reared in the Adelaide 
teuton from U. lugens and developed as a soliraiy 
hyperputasitoid through Euploctnts sp. and a\ a 
gregarious hyperparasitoid through D. eucalypn. 

Fulophidae (genus & species indei.) 
FIG. 38 

The two specimens reared as hyperpatasitoids of 
U, lugem through D. eucalypti in the Adelaide 
region could not be identified to genus due to the 
poor condition of ihe material. They are different 
from the other eulophids recorded here and can be 
distinguished by the dorsovent rally flattened body 
and distinctive colour pattern. 

Family fcupelmkiae 

A nan (at us sp. 
FIGS 29-31 
This species is easily separated from the other 
chaleidoids associated with U. lugew by its 
distinctive body shape and banded wing*. The genus 
is cosmopolitan and known mainly as parasitoids 
of the eggs of lepicloptcra and Hoteropiera, 

vctn; pm = poMina^uial vein; R. = oeth ^ = sternal vein; is = nir^l tatmentt 


\ D, AUSTIN & G. R. M I l-N 

Fi&K 2G-J5 26-27. h.urxtoma sp., y; 26, Dor^il view ol alittunk; 27, lore wing; 28-29, Lateral view of bodv: 28, 
Phroit'Ltlth sp,. s , 29, AtiifSiaiuh sp., •? : 30. Fore wine ol Antisiofus sjj„ y ; 31*33, Doisal view of alitiunk: 31, 
A'laswtita sp., V ; 32. bupelmidue, genus .& species indei., or; 33, Podiobius sp., 9; 34, fore tfjjig ol Pediuhhn sp., 
y. & Huh! let* ol Pwliohms s;p„ r r y Scales, FtgS 26, 27, 2'J, 34 and .35 = 250 giftf. Fig. 28 = 0.5 mm; Fig. 30 = 
20<» ;tiii, I'jfli 31-33 = 1 50 «m, Abbreviations: mp = mesnpleuron; pm = posimui^inaJ vein; KC = .sojinm; si = 
scutellum; W = Vtfe'W^I V <-'' J1 ; ,s ~ tur^-at >egmenis. 



fl ? hi • hlf f\ /ectni ^P- J- & Dorsal view of head and alit.uuk; 37. lore wine;; 3K; Dorsal view orulimink 
ulopnidae, genus & species mdcl., fl (stippling indicating colour pattern); 39-42, Wdimyumriia to.! W I ore 
T h,.K svmg, o-; 40, H.nd leg, 9t*l>? antenna; 42, ? antenna. Scales: Figs 36 and 37 = 250 „.n; hi » and 
39 = 150 [tm; Ffj: 41.) - 100 |/m; 1-igs 41 and 42 = 50 ^m (same scale line)- Abhrcviati 
vein; st = .>ugmal vein; is = tarsal segments. 

ions: pm 


although a few ate recorded as hyperparasitoids 
through braconids. Artastaius is represented in 
Australia by 40 described species (Boucek 1988), 
The species recorded here is represented by three 
specimens reared as a gregarious hyperparasitoid 
through C iirabae and £). eucalypti in the Adelaide 

Fupelmidae (genus & species indei.) 
FIG. 32 

A single male specimen reared as a hyperpara- 
sitoid of U, lugem through C. urabae in the 
Adelaide region could not be identified io genus, 
but it is noi Anastatus sp., from which ii can be 
separated by the characters in the key. 

Family Triehogrammatidae 

Trkhogramma sp. 
FIGS 39-42 
The members of this cosmopolitan genus are 


A. D. AUSTIN & G. |i. A! I FN 


Fit^s 41-49. 43-47. CetUradora sp.: 43, Fore win*, ,<; 44. Dorsal wl*w of whole ho<ty, ■?; 4\ J antenna; 46. er 
antenna; 47, Hind lee, V: 48. fifuitMiwil of ll'htt hernia lateralis. ■? (MacviuarO; 49, Dorsal \Un\ nf /-'vom/c/ /7</wtv/>_s 
Macquarl. "-> (stTpplmg showing colour pulieni in Figures 46 and 4/|. Scales: Fig. 43 - 200 ;iim Fig. 44 s 300 M m, 
Fi L -s4S-47 = a0jtw (sHmcsvjile tine for Figs 45 and 16); Fi^4fcanit49 = 1.0 mm. Abbreviafion: Is = tuisal ^lueuts. 

obligate, primary parasitoids of insect eggs, most 
frequently those of l.epidopiera. li is the only egg 
parasitoid of U. lugens so far recorded;, and it can 
be identified by the characters in the key, as well 
as its minute size, distinctive lore wing venation, 
fore wing setal paiLcru, and very narrow hind wing. 
This species is a solitary parasitoid known only from 
the Adelaide region. We have not. seen material of 
(he Inchograttima sp, reared from U. lu\>rm in the 
Murray Valley in N.S.W. (Campbell 1962), which 
may be tiie same species to that recorded here* 

Family Aphelmidae 

Centrodora sp. 
FIGS 4.V47 

This is a cosmopolitan genus of about 40 
described species (llayai 1983), mosl ol which are 
primary parasitoids of the eggs of Orthoptera and 
Homoptera, although at least one species is 
reported to be hyperparasitic (Gordh 1979; also see 
Viggiani 1984). The species recorded heie is an 
obligate, gregarious hypcrparasitoid of 11 lumens 

PARASiroms oi 0HAB4 LffGENS 


through C urahac or D eucalypti. Apart from the 
chatacters in the key and Hie life history stage 
attacked, Ccntrodom sp. can he separated from the 
other paiasitoids associated with U. lumens by us 
minute size, wing venation, and ovipositor which 
is more than half ihe length of the gaster (the 
ovipositor being MsmTicanily less thaii hall" Ihc 
gaslral length in Trkitaxramwa sp.). 


Family Tachhridae 

Wintlwmia lateralis IMacvjuan) 

FJG. 48 

Previously this species has been collected at 

various localities in all stales of Australia. It has 

been reaied from host species belonging to seven 

lepidopteran families, viz. Atctiidae, Noctuidac, 

Pieridae, Notodomidae, Nymphalidac, Saturnidac 

and Gcometridne tQosskey 1973; Cantrell 1986, 

19X9). The only record from U lugetis is from 

specimens in this study {3 specimens, Waite institute 

campus, J. Cobhinah, 1975, WAR!), lit toterelis 

oviposits onto the externa! surface of a hosl larva. 

After hatching the fly larva penettates the host larva 

and usually emerges from Ihc host pupa, although 

we were not abtc to confirm its biology m this study, 

Set Ctusskey (1973) and Cantrcll (1986, I9K*)) for 

additional taxonnmic information and list of 


txon.sia flaviceps Macquart 
FIG. 49 

This species has been recorded from all slates of 
Australia and the NT. (Cantrcll 1985), and has been 
reared from members Of nine lepidopteran families, 
W7. I.ymanlnidae, Anlhelidae, Pieridae, Agarts- 

9idac« Sphln&idac, Geometridae, Notodomidae, 
lawoeampidae and Noctuidac (Crosskey 1973; 
Canitell I9-S6). h oviposits onio the surface of U. 
luzem larvae After hatching the fly larva bunows 
into the host to feed and develop internally, finally 
emerging from laic larval instats fn pupate outside 
ihe dead host. The colour pattern on the abdomen 
and occiput of the head is the easiest way io dis- 
tinguish (his species from W. lateralis. SeeC'artliell 
(1985) for additional taxonomic information and 
list of synonyms 

Olher Parusifoids 

lYom U higetu in Queensland Brimhlecomhe 
(1962) reared two iehncumonoid species which have 
not been recorded in S.,, viz. Jnthufha sp. 
(Ichneumonidae) and Campyloneuro sp. (Bracon- 
ktac). Also Gauld (1^84) reports the following 
iehncumonids as having been reared from U fugertx 
Stiromesostenux spp., Campoplex sp. and 
Pristottterus _>p„ but again, they were not reared 
during this study and may not be found in S. Aust. 


Wc thank Ur Ian Naumann ( Australian Naiiunal 
Insect Collection), Mi Geoff Hollowav (Australian 
Museum) and Dt Bryan Canirell (Queensland 
Department of Primary Industries) for assistance 
with identifications: Mr Paul Daneerfield for the 
line drawings and scanning micrographs, and Dr 
Mike Keller lor reading a draft of ihe manuscript. 
This work was supported by a Commonwealth 
Postgraduate Research Award to GR A and a grant 
from the Australian Biological Resources Study 
participatory program to ADA. 


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hiaconids (Hymenopicrn) parasitic on Dlatrueu spp. 
(Lepidoptera; PyraliiJac). Bull. e/?/. firs 1% 131-144. 

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iurmiies, with a reclassified! Ion of specie*', ft AK 
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(Walk.}, ihc gum leal skcV-iom/er moth, with particular 
reference to the Fin\tlypius curnaMutenms Dchn. (river 

red gum) forests ol ihe Murrav V.iHey Region. t*wc t 

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Tor urowrh oi (he larvae ot Uruhu ttayth (W.ilk.). h»; 

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snpraspeeifie ia\aand laxouomJe andliofit i^tlfqnjcsi 

filtft. Br, Mux. nut. Hist, (Cnl.) 21, \-221 
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its parasites in Australia \./. J. Set. TechnoL (A). 21, 

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in the Hymenoptera. Proc. R. ent. Soc. Lond., Ser. A. 

43, 66-72. 
Galld, I. D. (1984) "An introduction to the 

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Hisloiv), London), 
Gordh, G. (1979) Supcrfamily Chalcidoidea. pp. 

743-1043, In Krombein, K. V., Hurd Jr, P. D.. Smith, 

D. R. & Burks, B. D. (Eds), "Catalog of Hymenoptera 

in America North of Mexico, I". (Smithsonian 

Institution Press, Washingion, D.C.). 
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lumens in Victoria. For. Tech. Paper, For. Common. Vic. 

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Hm^ris, R. A. (1979) A glossary of surface sculpturing, 

Calif. Dep. Food Agric. Bt4r, Ent, Occas. Pap. 28. 1-31. 
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(Hymenoptera) of the world. Syst. Ent. 8, 63-102. 
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paraphykticichneumotiid genus (Hymenoptera), and 

a revision of the Australian species. J. Nui Hist. 22, 

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Apante/es Foerstcr (Hymenoptera: Braconidae); a 

phvtoaenv and reclassification of Microgastrinae, Mem. 

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of Hymenoptera. Proc. ent. Soc. Wash. 88, 1-7. 
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Rev. Ent. 29, 257-276. 




byS. Barker* 


Astraeus meyricki Blackburn is resurrected from synonymy, redescribed and illustrated. Astraeus 
badeni van de Poll is redescribed and illustrated and the following five new species of Astraeus 
(sensu stricto) are described: A. aridus sp. nov., A. occidentalis sp. nov., A. princeps sp. nov., 
A. williamsi sp. nov., A. yarrattensis sp. nov. A revised key to Astraeus (s.s.) is provided. The egg 
of A. pygmaeus van de Poll is illustrated and some biological information given. Two new species 
of Stigmodera (Castiarina) are described: S. garnettensis sp. nov., S. wellsae sp. nov.; the first has 
sexually dimorphic colour pattern, rare in the sub-genus. 
KEYWORDS: New species, Astraeus, Stigmodera (Castiarina), Coleoptera, Buprestidae. 

pWWCtfclto Of rtd ftoytil SOCbty Vtf K 4Wf., H9H9), UJ(4| 185- Wi. 



by S. Barklr* 


BarMK. S. (I9B9) Contributions lo Ihc taxonomy of Australian Buptcsiidac ('. oleopteral: Not SpCEItl 
of mats und Stwtwderv (Catftarmaj and a key to .4,,\rwe/v.s (Xw, Tntns. N ,W . ,S »/*>/ 1 1.1(4), TK5-IV4. 
30 November 1989 

Astnwiis meyricki Blackburn is resurrected from synonymy, redeseribod and illuMrated. Uiruvu.s baitetit 
van de Poll is re-described and illustraJed and the following five new specie? ol Astraeus f&$ftft itrietQj 
are described: /f andus s-p. nOV r , A ocfhh'titahs Bp. nov . I- primvps -p. nov t willumtsi sp nov ., t. 
yarraftefSh sp. nov. A revised key to Astraeus fs.v.y is provided. I he egg of .1. py^maeus van de Poll is 
illustrated and some biological information given. Iwo new species ol Sn^Dhuh-ra (CaSttQflttu) *^. desirihrd 
S. gornettfUSiS sp, nov., S trW&tfff sp. nov,; the FJrsi ha? sexually dimorphic colour pattern, rate in lite 

Krv Words: New species, Aslrarus, Sttsmodera (Cus(tantni), Coleopiera, Bupresttdac 


The buprestid genus Asiraetis was revised by 
Barker (1975) and additional species described by 
Barker (1977). No further information has been 
published. Fresh material has now come to hand 
and live new species are described herein including 
one species from the Minders Ra., S. Aust., which 
was misidemified in the revision of Barker tl975). 
The separate status ol A. badenhww de Poll and 
A, meyricki Blackburn has been in doubt since they 
were described. A re-examination of the specimens 
available has convinced me that they are separate 
species. The key to Aaraeus fs.s.) given by Rarker 
(1975) requires updating to include three new species 
published by Barker (1977), five new species 
described herein and one species resurrected from 
synonymy. A new key is presented and new locality 
records of two species recorded. The egg of A. 
pygmaeus van de Poll is illustrated, the first such 
observation of an Astraeus species, together with 
biological information on A. pygmaeus. Two new 
species of Stigmodtra (Castiarina) are described 
and illustrated; one being unusual in showing sexual 
dimorphism in colour pattern, a rare occurrence in 
this sub-genus. 

Materials and Methods 

Male genitalia were prepared and displayed by 
the method described by Barker <I9K7). Specimens 
for S.E.M, were dehydrated prior to being coated 
with gold-palladium. Abbreviations used in the lest 
for museum and private collections following Wall 

(1979) arc: UMNH British Museum (Natural 
History), London; .IHQA Mr .1. Kasetipusch 
lunisfail; MNHN Museum National d'Histoire 
Naturellc, Paris; NMVA Museums of Victoria, 
Melbourne; SAMA South Ausirallan Museum. 
Adelaide; WAMA Western Australian Museum. 
Perth; GWNA Mr C. Williams, Laudsdownc; 
GWQA Mr G. Wood, Atherton 

Resurrection of Astracus rnryrickh 

Van de Poll (1892) considered ,4. meyricki 
Blackburn a synonym of A. hadeni van de Poll but 
Blackburn (1805) disagreed Barker (1975) followed 
van de Poll, I have re-examined the two male 
synlypes of A. meyricki held respectively in the 
BMNH and SAMA collections and compared them 
with specimens tn the SAMA identified by 
comparison with the female bolotypeof A. hadeni 
held by the MNHN. I he pronotum o\' A, hadeni 
is widest medially and diflers from A. mvynckt 
which is widest basally. Male genitalia of A, 
meyricki are broader than those of A. hadeni and 
are rounded more abruptly HI the apices (Tigs IC 
IP). I conclude thai A meyricki |f ie 2B) is j good 
species and hereby resurrect it from the synonymy 
ol' A. hadeni (Tig, 2A). Both species are tedesenbed 

AsiMtm iA*lr&eus) witttaMjl sp. nov. 
FIGS 1A, 20 

Hohavpc: Ef, Peninsular Rd, Laiisdownc, N.S.W , 
I0W19S7, G Williams SAMA I 21219. 

• Department of Zoologv Lfinvwsiiy ot Adelaide 
G.P.O. Bon 4% Adelaide, 5 AlKL SflOl 

A ffaiypz o , same data as uolotypc, SAMA 1 21220, 
Pararypey N . S.W ; 5 cr ct , Pen i nsu la r Rd . 






ll m m I 


Fig. I- Photomicrographs of male genitalia of the following Asirueus (Astraeus) species: A. Astraeus wi/liamsi sp. 
nov., B. A, aridus sp. nov., C. A. meyricki Blackburn, D. A. occidentals sp. nov, t E, A hadent van dc Poll, F. 
A. yarrattensis sp. nov,, and the following Stig/vodcra (Castiarina/ species: G. Stigmodera garnettensis sp. nov,, 
H. 5. valtist Dcuquet, I. S. welteae sp. nov. 

Lansdowne, 17.xi.1980, 23.X.1980, 30.X.1980, 
22.X.1982, 28.X.1988, G. Williams, OWN A. 

Colour: Head hiue-grcen or blue. Antennae, 
segments: 1-3 coppery; 4-11 blue. Pronotum blue 
laterally, purple medially. Elytra dark blue, each 
elytron with following yellow markings: basal spot; 
pre-medial fascia represented by mark over humeral 
fold and spot near suture; post-medial fascia 
touching margin but not reaching suture; small pit 
apical spot. Ventral surface dark blue. Legs: femora 
dark blue; tibiae mostly dark blue, distal tip 
testaceous; 1st tarsomere mostly testaceous, distal 
tip blue, 2nd tarsomere mostly testaceous, distal tip 
blue, 3-4 tarsomeres blue. Hairs silver. 

Shape and sculpture. Head closely punctured, 
without median keel, hairy. Pronotum closely 

punctured; laterally parallel-sided at base, rounded 
posteromedial^ to apex, hairy. Elytra costate, 
intervals flat and smooth each with basal row of 
punctures; laterally parallel-sided basally, rounded 
posteromedian^ and tapered to sharp marginal 
spine; sutural spine sharp, rounded inner margin, 
humeral fold moderately developed, angled (vide 
Barker 1975 Fig. IE). Ventral surface shallowly 
punctured, moderately hairy, hairs short. 

Size. Males, 7.5 ± 0.18 x 2.9 
Females, 7.9 x 3.2 mm (2). 

0.08 mm (6). 

Male genitalia: (Fig. 1A). Parameres parallel-sided 
from basal piece, widened medially, tapered, 
rounded and tapered to apex. Apophysis of basal 
piece broad, rounded apically. 



Fig. 2. Habitus illustrations of the following species: A. Astraeus badeni van de Poll, B. A. 
meyricki Blackburn, C. A. aridus sp. nov., D. A. wi/fiatmi sp. nov., Li. A* occidentalis 
sp. nov., F. A. yarratiensis sp. nov., G. A. princeps sp. nov., H. Stigmodera wettsae sp. nov. 



Remarks: Named after Mr G Williams, Lativ 

4iftfietii MtfyafosJ >a*rsr*cnsis sp in-v 

MCiS IP, 2f 

Astmeus (Astrueus) aridus sp. nov, 

Astrueus haden't: Barker 1975, p 131. 

Holotype: or. Puttapa Gap. 21 km S Copley, S. 

Ausi., Flinders Ra., 2I.X.1971. S. Parker, SAMA I 


Allotvpe: 9 , same data as holorype, SAMA I 2)224- 

Paratopes: S. Aust.: 6 <y o\ 8 99, same data 312 
Allotvpe ?i 10 km SSW Laurieton. N&W. a holotypc SAMA; 8 t?(y t 6 9 9. Dema Pass. 
&xr.l*80< & Williams, SAMA I 21222. SSXlBWi A*. Mcfurland, SAMA, 

Hokwp*- -. Yarrau State Forest, NF Winghum, 
N.S.W., tl.xiinst, a William*. SAMA I 21221, 

Puratypes: N.S.W.: t -<*. *amc data as holotypc, 
CWV'V A :• v. Strati Siatt Forest, NE Wingham, 
29.x.l'W. 16.\i.l ( v82, O, Williams GWNA. I o\ 
Bom Bom Stole loresi.SF Grafton* l4.tii.T984, 
H t/haitis, GWNA; 1 w Mitchells' Maud, (Factory 
Rd)> U Taree, 9.xi.l988, C. William^ GWNA. 

C o/bWA Male. Head blue-green, dark blue basally 
With purple reflections. Antennae, segments: 1-3 
coppery; 4-11 blue. Pronotum dark blue with 
put pie reflections, blue at lateral and basal margins. 
Female. He;id and pronotum datk blue, rest as in 
male, Elytra dark blue with following yellow 
markings on each elytron: large basal spot; pre- 
medial fascia covering humeral fold bul not 

teaching suture; poRt-medial fascia covering margin 

but not reaching suture Ventral surface blue. Legs: 
femora blue; tibiae blue at proximal end, rest 
testaceous; Isl tarsoiueie testaceous; 2-4 tartOIBCWS 
blue, Hairs silver, 

Shape and sculpture: Head closely punctured, 
glabrous median keel, hairy, Pronotum closely 
punctured; laterally parallel-sided basally, rounded 
postcromedially, tapered to apex; median impressed 
line from middle to apical margin, laterally hairy. 
Elvira costale, intervals flat, basally wrinkled, each 
with row of punctations; laterally tapered from base, 
rounded postcromedially, tapered lo sharp marginal 
spine; shatp surural spine, rounded inner margin; 
humeral fold well developed and angled. Ventral 
SUI face closely punctured, moderately liauy, hairs 
medium length. 

Si:c: Males, 6.9 x 2.9 mm (6). Females. 7 4 \ VI 
mm (2). 

Male genitalia: [ftfc. IH. Parameres angled 
outwards from basal piece, more or lev* parallel- 
sided, rounded w apes Apophysis of basal piece 
brOad, rounded apieally. 

Remarks. Named aftei the ivpe locality. 

Colour: Head, antennae, pronotum black with blue 
reflections. Elytra black with hlue reflections and 
following yellow markings: basal spot; fascia 
covering humeral fold, not reaching suture, concave 
anteriorly, often represented by two spots; short 
post-medial fascia touching, margin bul not suture, 
cofn:ave posteriorly Ventral surface and legs black 
with purple reflections. Hairs silver. 

Shape and sculpture: Head closely punctured, no 
median keel, hairy. Pronotum closely punctured; 
laterally rounded and narrowed from base to apex; 
hairy. Elytra costate, intervals flat, each with row 
Of punctures; laterally parallel-sided basally, 
rounded posteromedially lo marginal spine; both 
spines sharp, sututal spine with rounded innei 
margin, humeral fold poorly developed, slightly 
anpled, Ventral surface shailowly punctured, 
moderately hairy, hairs medium length. 

Size: Males, 8.9 ± 0.10 x JJ t 0.07 mm (15). 
Females. 9.1 + 0.14 x 3.6 f 0.07 mm (15). 

Male genitalia: (Fig. IB). Parameres angled 
outwards I torn basal piece, rounded anteromedially 
to apex. Apophysis of basal piece medium width, 
rounded apically. 

Remarks: All specimens were collected on the leaves 
and stems of Melaleuca slomerata F. Muell. from 
Iht* same locality. Oerna Pass is a synonym ol 
Pultapa Gap. The specific name is derived from 
aridus L., dry. 

Asfnjeus (Astracus) hadeni van de Poll 

AslniHis Intihtu van e|c Poll 1X89: 84, 91, 94, pi. 2, fics 
7, 7i<. Blftvtfcurn I891{ 496; Kcrrcmans 1892: 101, van de 
Poll 1892: 67; Blackburn 1893: 45, 46; Kerremans 1903 
I4K; Carter 1929 282; Ohcnherger 1930: 365; Barker 1975: 

Astrueus budeni disjunctw; Obcnbcrser 1928: 204; 1930: 
365; Marker 1975: 131. 

Asiraeus mevricki Blackburn 1890: 1256, 1257; van tie 
Pall 1892. 67; kerremans 1892: 101; Rer remans 1903: 148. 
Cartel 1929: 282; Ohcnherger 1930; 365: Barker J 975: 131. 

Tvpe: Holorype 9, Cawler, MNHN. 


Colour; Head, antennae, pronotum black with blue 
and purple tcfleetions. Elytra black with purple 
icfleclions and following yellow markings on each 
elytron: basal spot; prc-tnedial fascia from margin 
over humeral callus, not reaching suture, concave 
anteriorly; shori post-medial fascia from margin not 
reaching suture, concave posteriorly; small pic 
apical spot. Ventral surface black with biun/eand 
purple reflections. Legs black with blue and puipie 
reflections. Hairs silver. 

Shape ami sculpture: Head closely punctured, no 
median keel, hairy. Pronotum closely punctured; 
laterally parallci-sided nasally, rounded, indented 
to apex, hairy. Elyira costate, intervals flat each wilh 
row of punctures; laterally paiallel-sidcd basally, 
rounded posteromedial!); abruptly tapered to sharp 
tiinr i^in;H spine; sharp surural spine, rounded inner 
margin, humeral fold moderately developed, angled. 
Vential surface shallowly punctured, njodcrately 
hairy, hairs long, 

Size; Males, 7,9 x 3.4 mm (6). Females, S.4 x 3.6 
mm (8). 

Mate genitalia: (Fig, IE). Parameres angled 
outward* from basal piece for most of length, 
rounded to apex. Apophysis of basal piece medium 
width, rounded apjcally. 

Distribution: S. Aust.: Gawlcr. Morgan, Murray 
Bridge, Nuriootpa. Vie.; Inglewood. W.A.: Marvel 
loch, Southern Cross. 

Remarks: A, hadeni differs from congeners in the 
bulbous appearance of the pronoium and the more 
abruptly tapering apices of rhc elytia. It is a short 
broad species. A specimen in the SAMA collected 
by Tcpperai NutiooLpa, S. Aust. has 'on Cathtris' 
on ftic label, ihe only specimen seen with a recorded 
h»iM plant. 

Astraens (Astravus) meyriekt ttlneMnKn 
TIGS 1C, lb 

Auracus- nwvtteki HkKkbwn, I84(): 1256. 12s?; NPJtJ fa 

Type: Syntype i/, no data, BMW I; Byniype i ?, W. 

Australia, SAMA. 

Colour, Head, antennae, prunoinm black wjth 
purple relfcvtions, Hlyrra black with following 
yclluw marks on each elytron: small basal spot; pre- 
medial fascia covering humeral lold, not reaching; 

suiure; post-medial laseu not reaching maigm or 
suture. Ventral surface black wilh purple reflections. 
Legs dark brown with purple reflections. Hair* 

Shape and sculpture; Head closely punctured, 
shallow basal median sulcus, without median keel, 
hairy. Pronotum closely punctured, wilh prominent 
basal fovea, median glabrous line trorn base Lo 
middle; laterally rounded from base to apex, Flytra 
costate, intervals flat, each wilh row of punctures 
hue-rally tapered I mm base, rounded posteromedi- 
al^ 1 then tapered to sharp marginal spine; sharp 
sulurat spine, rounded inner margin; humeral fold 
very poorly developed, very slightly angled. Ventral 
surface shallowly punctured, moderately hairy, flairs 
medium length. 

Size; Male, 10.6 % J.5 mm (2). 

Male genitalia: (Pig. IC). Paiameres angled 
outwards from basal piece, rounded to apex. 
Apophysis of basal piece medium widih, rounded 


Remarks There is a female specimen lodged in thc 
SAMA possibly belonging to this species which J 
identified a.s/4. badeni (J3arker 1975 p< 13.1). It wa* 
collected IS km SW Three Springs, W.A,, on 
Dryundra cirisoides\ 8..\i.l968, N McFurland, 

Astraeus (Astmrus > princess sp. tine 
FIG. 2G 

Hoinlype; <?. Piince of' Wales Island, Qld, Feb 
I939, SAMA I 21225 

faratype: Old: 1 e„ some data as holotype SAMA. 

Colour; Head black with blue reflections and yellow 
frontal spot. Antennae black with blue reflections. 
Pronotum black with blue reflections and large 
lateral yellow marks. Flytra black with blue 
reflections and following vellow marks on each 
elytron; broad prc-medial fascia covering margin 
but not reaching suture, obliquely angled anteriorly; 
broad poM-medial fascia covering margin bur not 
teaching suture. Ventral surface dark blue. legs red 
brown, Hairs silver. 

Shape and sculpture: Head shallowly punctured, no 
median keel, few short hairs. Pronotum shallowly 
punctured, laterally rounded from base to apex, 
widest medially, few shon hairs. Elytra costate, 
intervals Hal, each with row oi punctures, suiface 
wrinkled nasally; laterally tapered trorn bass; 
rounded postcmmedialiy, tapered to sharp marginal 



spine; sharp sulural spine, inner margin straight and 
turned upward, with extended marginal interval 
between spine**; humeiaJ fold absent. Ventral surface 
shdllmvly punctured, moderately hairy, hairs 
medium length. 

Size: Females, 7,9 x 3.4 mm (2). 

Remarks: Both specimens were given In Mr S. 
Watkins by Rohert G. Wind, insect supply dealer 
and resident of Monterey, California. His Australian 
wife had been stationed on Prince of Wales Island 
prior to World War I I and presumably she was the 
collector. They are unique to the sub-genus due to 
the coloration on the head and pronotum. The 
dorsal coloration may have been pink or red in life 
and now faded. The name is derived from princeps 
L, leader. 

Astmcus (Astraeus) occidentals sj> iiov 
HGS ID, 2b 

floiotype: o\ 99 km S Overland Road house, 
Carnarvon Hway, W,A, ( 25.ix.i980; on Casuorina 
(Jk'ls(urta % S. Barker, SAMA, I 21226. 

Allotype: v 1 . same data as holotype, SAMA. 1 

Paralypes: W.A.; 2 u-o*, j £} WAMA, SAMA. 

Colour Head, antennae, pronotum coppery-bronze. 
Elytra black with blue and purple reflections and 
loltowine yellow markings on each elytron: basal 
spot; pre-medial fascia covering humeral fold, 
touching basal margin but not reaching suture, 
concave anteriorly; medial spot on margin and one 
near suiuie in .some specimens; large post-medial 
spot near margin; pre-aptcat spot near suture. I lairs 

Shape and sculpture: Rend closely punctured, I bin 
medial impressed line from base, continued as short 
jjlabunis median keel al apev hairy. Pronotum 
closely punctured; prominent basal fovea projecting 
anteriorly for short distance as impressed lino, 
anlerior margin projecting medially; laterally 
rounded and narrowed from base to apex, hairy, 
hlytra coslate, mieivals flat, each with iow of 
punctures; laterally parallel-sided basally, rounded 
medially and narrowed to small marginal spine; 
.dwirp.suturul spine, rounded inner margin, humeral 
fold poorly developed, slightly angled. Veutial 
surface shallowlv punctured, moderately baity, hairs 

Size: Males, 11,2 \AA mm 0). Females, 11,8 x 4.6 
mm (2). 

Male genitalia: (Fig ID) Parametes angled 
outwards from basal piece, slightly rounded 
medially, rounded nbruptly to apex. Apophysis of 
basal piece broad, rounded apically, 

Remarks: The name is derived from occidenfuti$\., t 

Key to the species of Astraeus (seam uric(o) 

Head with median keel 

Head without median keel, , 

Hans silver. 

Hairs yellow 

Pari or all of anterior ventral surface red-brown 

.,, 4 

None of anierioi ventral surface red-brown 7 

Gular. presternum, ruesu- and tneiasternum, coxae 

2-3, abdominal segment I red-brown , , 

c . A. hakeri Barker 

Less O.f ventral surface red-biown. . . , , ; . 3 

Prostcrnum, coxae ted-brown , ( , 

, . , .A, mimaus Barker 

Red brown area on either side flf prosteniul process 







Shorter than 7,5 nun; humeral told well developed, 

ucutelv angled 4. fraserietws Barket 

Longer thnn 7.5 mm; humeral fold moderately 

developed, angled A. ob$twns Barket 

Most or part of leg lestaeeous.. 8 

None of leg le.siaccous- 13 

Legs 1-2 testaceous except Tor outer margin ol lemoia, 

leg 3 testaceous except for lemur , , 

, , A- diliaipes van de Poll 

1 ess of leg than above testaceous 

Tibiae, myomeres 1-2 testaceous , , 

„. r ._..,, , A. smyrhi Barker 

Less of leg than ahove testaceous. . . . 10 

Tibiae testaceous, distal tips blue, tarsomcre I 

testaceous .-1. yitrnatensis sp. no\. 

Less of leg than above testaceous it 

Distal tips of tibiae testaceous* tarsomeres 1-2 
testaceous, dtsuil tips blue. , ./L wifltamsi sp. nov 

Less of U>* than above u^tacc-ous \1 

lips of tibiae and larsomere I testaceous I? 

TiMSOuieie I loia.A'Oiiv , , , 

A. mtjawiMt'o< , > Baikrt 

Usually shorter than 7 mm 

A. {tyxmaeas van do Poll 

Usually longer than 7 mm , , 14 

Humeral told well developed, acutely angled. , , , . 

._,,..,,,, . , A, trwstersi MacLeav 

Humeral fold well developed, angled 

A, sammwtli Saunders 

Humeral told uiudc-iatelv developed, angler). . . . U» 
Humeral fold poorly developed, slightly aueleiL 




16 Head green and coppery purple or blue-green. - 17 
Head black 18 

17. Head, pronoium green and coppery purple , 

- i> — , i . . -A infricatus Carter 

Head blue-green, pronoium with medial, cordiform 

purple mark, anieriorly green, laterally blue 

- . A, blackdownsensis Barker 

18. Broad, rounded species A. xfobosus Barker 

Elongate species 4. wat&ani Barker 

19. Body blue; elytra with 2 yellow fascia 

.... i , , , .A. fraWrculux van de Poll 

Body black or coppery- bronze -20 

20. Body black; elytra with numerous yellow spots. .. 

. . .A. crassus van de Poll 

Body coppery-bronze; each elytron with yellow fascia 
and 4-S yellow spots A. 'uvultntalis sp. nov. 

21. Klylra with 3 yellow fascia, red areas. 

A, major Blackburn 

Elytra with 2 yellow fascia, no red 

4. fiuvnrchi\ { Thomson) 

22. Body elongate and cylindrical 23 

Body tear-drop shaped 24 

23. Pronoium conically elevated medially. 

.4. profhoracicus van de PolJ 

Pronoium convex medially _ . 

'1. elongatus van de Poll 

24. Sulurul spine with rounded inner margin ... 25 
Sutural spine with .straight inner maigin 37 

25. Legs red-brown colour 26 

1 CftS other than red-brown colour 27 

26. tlytra marked with spots and fascia , 

A, mavmUiani Barker 

F.K-h elytron with 2 viltae A. vittatus van dt Poll 

27. Head, pronotum, legs metallic brown or bronze. 

A. ftavopictus C & G 

Head, pronotum, legs other than brown or bron/c 


2H. Humeral lold well developed, angled, ...... .29 

Humeral fold moderately or poorly developed. .30 

29. Head bl»ck or coppery purple; ventral SUffnCC 

coppery-purple L .4. uriamsi Barker 

Head blue oi green; vcnlral surface blue-green... 

A. simulator van de Poll 

30. Humeral fold modeiatcly developed, angled, ,31 
Humeral fold poorlv developed, slightly angled. 


31. Pronotum laterally rounded from base to apex.,. 

-4. crockerae Barkei 

Pronoium parallel-sided at base, rounded, indented 
io apex , , ,4. batleni van de Poll 

32- Basal spot touching basal margin Of elytron 

4. cartiahyi Barker 

Basal spot not touching basal margin of elylron 
n 33 

33. Elytron with 2 spots and 2 fascia or 4 spots and a 

fascia , 34 

Fly Iron with 6 spots and a fascia, 7 spots or 8 spots 

34. Head pronoium and elytra black . , 

.A. miyrickl Blackburn 

Head pronoium and elytra black, with blue And 
purple reflections A, aridus sp. nov. 

35. Each elytron with ft spots and a tascia or « spots 

M i A\ fanSQHt van dc Poll 

Each BlyiKKl with 7 *:poN 3ft 

36. Pronotum parallel-sided I mm base to middle, strongly 
rounded and narrowed lo apex; dorsally convex in 

lateral profile 4. ohenhun v;m dc Poll 

Pronoium gradually rounded laterally, narrowed from 
base io apgfcj ilorsally flattened in lateral profile 

- 4. tvrreri Barker 

!7, Head with frontal spot . . ,A. prinveps sp. nov. 
Ik-ad ttrfrtlOQfl, frontal spoi , , .38 

38. Head with shallow median sulcus. 

--- A. gtoerlingi Barker 

Hfiad -a nil deep median sulcus 39 

39, Head with basal median sulcus; pronoium laterally 
inflarcd, medial oval patch of hexagonal cells. 

.4. cViJrwu.s Kci remans 

Head with apical median sulcus; pronoium laterally 

rounded, no medial hexagonal cells 

- I. caletJonkvx l-auvcl 

New locality records 

A single female specimen of A irregularis vai) 
dc Poll was collected by Mr il F. Adams and myself 
at Blackdown Tableland, Qld, on 17.1.1979 on i he- 
foliage of Custtarina inophloia. This is a common 
W.A. species recorded from S. Ausi. by Barter 
(1977). In the revision of Astraeus (Barker 1977) I 
did not record A, jansoni van de Poll outside of 
S. Aust, except for a doubtful Queensland record- 
Carter (1929) listed the species as occuirilig in the 
Blue Mrs. I have examined Garter specimens in the 
SAMA collection from the Blue Mts and more 
recent material collected by Mr S. VVaikms on 

r'i.u. 3. S.L.M. micto^mh ul the chorion ..< \yri 
pyxtnaiWi \an dr Poll 



Callitris muelleri at Wentworth Falls and Kings 
Tableland in the Blue Mts and confirm them to be 
A. jansoni. 

Egg of A. pygmaeus van de Poll 

A female specimen of Astraeus pygmaeus 
captured by me 24 km east of Armidale, N.S.W., 
on 19.xii.1989, laid 13 eggs onto the cardboard label 
in the vial in which it was placed. These hatched 
approximately 28 days later. The hard sculptured 
chorion of an egg from which a larvae hatched is 
illustrated (Fig. 3). Crowson (1981) indicated that 
most beetles have smooth eggs with a soft surface. 
A micropyle cannot be distinuished, but a row of 
tubercules, each pierced by an aeropyle, is clearly 
visible around the margin (Fig. 4). The eggs are oval 
with a flat ventral surface. The larvae emerged 
through the ventral surface and burrowed directly 
into the label. The chorions adhered to the label 
and covered each individual gallery. Presumably the 
same happens in the field situation. These are the 
first eggs of Australian buprestids that I have seen. 

Fig. 4. S.E.M. micrograph of aeropyles (arrowed) on 
chorion of Astraeus pygmaeus. 

Stigmodem (Castiarina) garnettensis sp. nov. 
FIGS 1G,5 

Holotype: o\ Mt Garnett, Qld, 24.ii.1989, G Wood, 
I 21228 SAMA. 

Allotype: 9, Mt Garnett, Qld, 24.ii.1989, I 
Hasenpusch, I 21229 SAMA. 

Paratypes: Qld: 1 9, Mt Molloy, 9.H.1987, G. Wood, 
GWQA; 2 9 9, Kuranda, 6/8. i. 1989, /. 
Hasenpusch, JHQA; 1 9, Mt Garnett, 20.ii.1989, 
G. Wood, GWQA; 1 tf, Mt Garnett, 21.ii.1989, 1 
Hasenpusch, JHQA; 2 o- o\ 3 9 9, 22.ii.1989, J. 
Hasenpusch, JHQA; 5 o* a\ 2 9 9 » same data as 
allotype, JHQA; 5 co", 6 9 9, Mt Garnett, 
21/22.ii.l989, G. Wood, GWQA; 10 cr cr, same data 
as holotype, GWQA; 1 9, Mt Garnett, 26.ii.1989, 
G. Wood, GWQA. 

Colour: Male (Fig. 5A). Head, antennae and 
pronotum coppery-purple. Scutellum green with 
coppery-purple reflections. Elytra yellow with 
following markings either black with coppery- 
purple reflections, or dark blue with green 
reflections: variable basal mark, continuous with 
basal margin, not reaching lateral margins, reduced 
to basal margin and extension along suture in one 
specimen; post-medial fascia reaching margin; 
apical mark. Ventral surface coppery purple. Legs 
green with coppery-purple reflections. Hairs silver. 
Female (Fig. 5B). Head, antennae and pronotum 
green with yellow reflections, occasionally head and 
pronotum coppery-bronze. Scutellum green. Elytra 
yellow with following dark blue markings: broad 
basal fascia; broad post-medial fascia; basal mark. 
Ventral surface green with yellow reflections. Legs 
blue-green. Hairs silver. 

Shape and Sculpture: Head closely punctured, 
median sulcus, short muzzle. Antennae, segments: 
1-3 obconic; 4-11 toothed. Pronotum closely 
punctured, narrow basal fovea extending forwards 
to middle as glabrous line, basal notches on each 
side more marginal than medial; apical margin 
projecting medially, basal margin barely bisinuate; 
laterally angled outwards from base, rounded one 
third of distance from base, bulbous, tapered to 
apex. Scutellum scutiform, punctured, flat. Elytra 
punctate-striate, intervals convex, punctured; 
laterally angled out from base, rounded at humeral 
callus, concave, rounded posteromedially, abruptly 
rounded to spineless apex; apices diverging, apical 
margin subserrate. Ventral surface with shallow 
punctures, edges of abdominal segments glabrous, 
moderately hairy, hairs medium length. S 7 rounded 
in both sexes. 

Size: Males, 14.2 ± 0.15 x 5.8 ± 0.07 mm (24). 
Females, 15.4 ± 0.27 x 6.6 ± 0.12 mm (17). 

Male genitalia: (Fig. 1G). Short and broad. 
Parameres angled outwards from basal piece, 
rounded posteromedially then parallel-sided, 
rounded to apex. Apophysis of basal piece medium 



/ mm ^ 

v\m } 1 1 i i 


Fig. 5. Habitus illustrations, of the following species: A. Stigmadera xarnettensis sp. nov, mule, B. S'. 
gurnet tensis sp. nov. female. 

width, tapered to rounded apex. Medium lobe 
pointed* sides obtusely angled away. 

Remarks: This is a member of the £ omenta 
Saunders species group as well as 5. flava Saunders, 
S. ovata Barker, S. pal/as Blackburn and S. vallisi 
Deuquct. Like S. vallisi the sexes are dimorphic and 
can be separated on colour and pattern differences. 
In the male genitalia of 5. vallisi (Fig. 1H) the 
apophysis of the basal piece is narrower. The 
specific name is derived from the type locality. 

Stigmodem (Castiarina) wellsae sp. nov. 

Holotype: cr, Portland, Vic, 21.xi.1944, E. Smith, 


Allotype: 9, 20 km N iMoe, Vic, 8.xi.l979, .4. 
Wells, SAMA 1 21230. 

Paratopes: Vic: 1 cr, 1 9, Emerald, 3.\ii.l907, 
Jarvis, NMVA, SAMA; 3 cro\ Grampians, Nov. 

1950, R Given, NMVA; 4 a cr, Portland, 6.xii.l918, 
NMVA; 1 o\ Fermree Gully, 30.x-.1919, NMVA; I 
cr, Portland, NMVA; 1 cr, I 9, Taruna', Croydon, 
S. W. Fulton, NMVA; 1 9, Bclgrave, 19.xi.I946, E 
E. Wilson, NMVA; 1 9> Healsville, Nov. 1916, 
NMVA; 1 9. Monbulk, I2.xi.l902, Jarvis, NMVA; 
1 9, Boulora, 4.xii.I954, A. I . Brown, NMVA; 1 
9 .locality indecipherable, 14.xi.I903, NMVA; 1 cr, 
4 9 9 . no data, NMVA; 4 cr cr, 5 9 9 , Mt Rosea, 
Grampians, 15.xi.l950, G. F. Gross, SAMA; 1 cr. 
Hall's Gap, Grampians, 14. xi. 1950, LeSouef, 

Colour: Head and antennae blue wilh bronze 
reflections. Pronotum blue with bronze reflections 
medially, yellow laterally. Scuiellum blue with 
bronze reflections. Elytra yellow with following blue 
markings with blue-green reflections: very broad 
basal mark only touching margin at humeral callus; 
broad post-medial fascia; pre-apical spade-shaped 
mark, reduced in holotype and allotype to short 
horizontal bar, all marks connected along suture 
to apex. Hairs silver. 



Shape and Sculpture: Head closely punctured, 
median sulcus, short milZZle. Antennae, segments: 
1-3 obeonic; 4-U toothed. I'tonotum closely 
punctured, narrow basal fovea extending anteriorly 
to apical margin as impressed line, shallow fovea 
ai each basal angle: apical margin projecting 
medially, basal margin barely bisinuate; laterally 
rounded from base to apex, widest medially, 
Seulellum scutifoim, lew punctures, flat. Elytra 
punctate-striatc, intervals convex, punctured; 
laterally angled out from base, rounded at humeral 
callus, concave, rounded posteromedial!) and 
narrowed to spineless apex; apices hardly diverging, 
apical margin rough. Ventral surface with shallow 
punctures, edges of abdominal segments glabrous, 
moderately hairy, hairs short. S 7 ; male truncate; 
female rounded. 

Size: Males, 10.1 ± 0.18 x 4.0 i 0.09 mm (19). 
Females, 10.5 ± 0.19x4.1 ( 0.10 mm (16). 

Male genitalia: (Fig. II) Short. Parameres parallel- 
sided nasally, rounded posteromedially, parallel- 
sided, rounded apically. Median lobe sharp, sides 

acutely angled away, Apophysis of basal piece 
medium-width, tapered, flattened apically. 

Remarks: Member of the S. distinguenda Saunders 
species group together with S. arida Barker, S. 
blavkdownensis Barker. S. garrawillae Carter, S. 
grata Saunders, S. milituhs Carter, S. subgrata 
Blackburn, S, supergrata Barker, S. watkinsi Barken 
It is closest to S. watkinsi but differs in having: blue 
markings on the dorsal surface whereas those in S. 
watkinsi arc green, a different distribution; different 
male genitalia (Barker 1988; Fig. \E). Specimens 
in the NMVA had been misidentified as S. grata 
Saunders. Named after Dr Alice Wells. 


I thank the following for their assistance: Miss 
C. M. H. von Hayek, BMNH; Dr A. Neboiss, 
NMVA; Mr G. G. Burns, Mornington; Mr .?. 
Hasenpuseh, Innistail; Mr S. Walkins, Caparra; Mr 
G. Williams, I-ansdowne; Mr G. Wood, Atherton; 
Mr P. G. Kempster, Mr C. Miller, Ms H. 
Vanderwoude,Dr A. Wells, Mr D. J, Williams, 
Department of Zoology. University of Adelaide. 


B/YRKER, S. (1975) Revision of the Genus A&UWUS'ortc & Gorv (Coleoptera: Buprcstidac). Trans, R. 

Soc, S. Atist. 99, 105-141. 
____ (|y77) Astraeus (Coleoptera: Buprcstidae): A 

description nt three new species and new locality 

records, tbid. 101. U-14, 
(I9S8) Contributions to the taxonomy of 

Stigmodera (Costiarma) (Coleoptera: Bupresiidae). 

Ibid. 112. 133-142. 
Blackul'Kn, I. (1890) Notes on Australian Coleoptera. 

with descriptions ot new species, part v. Proc, Linn. 

Soc. N.SM' 4. 1247-1276. 

(1891) Notes on Australian Coleoptera, with 

descriptions ol new spreics, fhirf, 5, 479-550, 
(1895) further notes on Ausimlian Coleoptera, 

with descriptions of new genera and species. Trans, R. 

Soc & AuSt 19, 27-60. 
Cakiik, H. .1. (1929) A cheek list of the Australian 

Buprcstidac. AuaL Zooi 5. 265-304. 

Ckuw-iin, R. A. (1981) The biology of the Coleoptera. 

(Academic Press, London.) 
Kf.RHEMANS, C, (1892) Catalogue wnonvmiquc dc* 

Bupreslidei de'erits de 1758 & 1890. Mem. Soc. em. Befe 

I, 1-304. 
(1903) Coleoptera; Scmcortiia Fam. RupreMidae. 

Genera Insectorura 12. 1-338. (P. Wytsman, Bruxellcs.) 
OBBNBEKGER ; J. (1928) Opuscula Buprestologica I. 

Beitrage /ur Kcnninis der Bupreitidcn (Col,), Arch. 

NofUfgeSCK 92, 113-224. 
(1930) Buprcstidac II. Coleoptorum Catalogue 

XII: 213-568. (W. Junk Berlin, Den Haag.) 
van or Poj i , J. R, H. N. (1889) Monographical essay on 

ihe Australian Buprestid seam Astraeus C. et Ci. Tjschr 

v. Em. 32, 79-110. 

(1892) Note SUT quelques c$pftcjcf UVt \tracu\. Ibid. 

36, 67-68 
Watt. J. C. (1979) Abbreviations for entomological 

collections N. Z. /.oot. 6. 519-520. 


by Richard J. F. Jenkins* & PamHasenohr 


An Early Cambrian trilobite from the Normanville Group in the Sellicks Hill area, Fleurieu 
Peninsula, South Australia is here identified as Ivshiniellus briandailyi sp. nov. The trilobites occur 
in 'black shale' in close stratigraphic proximity to a bedding-plane surface showing trails made by 
large trilobites. The environment was below storm-wave base and the traces indicate that the black 
shale deposited in water sufficiently well oxygenated to support large metazoans with a presumed 
oxidative metabolism. Black coloration in shales may not be taken as a necessary indication of 
anoxic bottom waters. 

KEY WORDS: new species, conocoryphid trilobite, trace fossils, black shale, Cambrian, South 

transactions 4/' the Royal Society of S. Aust., (1989), 113(4) 195 201 


by Richard I F. .if.nkins* & Pam Hasfnohr 


Jlnmns, R. J. L & Hasinohk, P. (1989) Trilobires and their [fails m a black shale; Early Cumbrian of 
ihe Hcui'ieu Peninsula, South Australia. Trans. R, Sac, S. Aust. 113(4), 195-203, ftfl NcHeuibei, 1089 

An Early Cambrian rrilobite I'roui the NortofltlVtUe Group in (be Selheks HJJI area, Heuricu Peninsula, 
South Australia is here identified as Ivslumettus hriamJailyi Sp. nov. The trilobites occur in 'black shale* 
in close straligraphic proximity to a bedding-plane surface showing Irails made by large nilobiies. Ihe 
environment was below uorm-wavc base and the traces indicate thai the black shale deposited in water 
Sufficiently well oxygenated to support large metazoans with u presumed oxidative metabolism. Black 
coloration in shales may not be laken ax a necessary indication of anoxic bottom waters. 

Ktv Words new species, conocoryphid tritobite, trace lossils, black shale, Cambrian, South Australia. 


Jago et ah (1984) reported the first discovery of 
trilobite remains in the Early Cambrian Norman- 
vilte Group on Fleurieu Peninsula, South Australia. 
They described two specimens of a conocoryphid 
trilobite from the Heathcrdale Shale in the Sellicks 
Hill area (Fig. 1), bul considered the material so 
poorly preserved as to preclude formal description. 
While exploring the same site during late 1984, one 

* Department of Geology and Geophysics, University of 

Adelaide, GPA BOX 498, S. Aust. 5001. 

of us (P.H.) discovered a spectacular trace fossil 
made by an arthropod. Furlher searching revealed 
two more body fossil remains of trilobitcs. This 
paper reports these new discoveries. 

Of the new trilobite remains* one shows the 
cephalon and anterior thorax and the other Is a 
fragmentaty thorax. They were discovered on 
surfaces that had split open to the weather, and the 
very incomplete specimen is preserved essentially 
as a ferru^inised film. The more complete individ- 
ual, an external mould, resembles specimen SAM 
P24321 described by JagO « al. (1984) and provides 


ia 30 JO U) -rO 

<#t I cc tan 

.\ Hang it* o iswn i i ■■., 

i .;■■ i ■ Brt na 
iii n-.i.-..v.,jiii, Otvjirp 
| I 3 AtielfliRMr frtawnw i -. 

, ..I., ,.5, I ... , .-,, 

lu- I. I Ocality map showing find ot trilobitcs and (race I'ossiU ai Sellicks Hill, and the placement of this tfMwcri 
in relation to the framework ot the Adelaide fold [Jolt tuu\ adjacent loreland basin- 



Fjg, 2A-C. /vshinieltttf hriandaUyi nov, sp.: A, holotypcSAM P24321, with anterior glabella relatively well preserved 
and composite mould of hypostome present, \3; H, C, para type SAM P24844, B, external mould and C, silicon 
rubber cast of individual made from mould, both xl.5. D. conocoryphid gen. et sp. indel, SAM P24845, fragment 
photographed under a film of water, xl.5. 



sufrtcieni additional morphological information to 
justify formal description of the taxon. 

The arthropod traces in near association with (he 
body fossil specimens of rhe rrilobites arc or especial 
interest as the host rock represents a 'black stinlc" 
faciei Thus Hie (races provide an unusual 
palaenbioloeical perspective on this interesting find. 

The studied specimens are registered in the 
Palaeontology Collection of the South Ausiiahan 
Museum, Adelaide (SAM). 


Siipoifamil) CONOCORYPHACi-.A Augeliti. 1854 
i-amily CON'OCOKYPHlDAfc Angelin, 1$5<1 

Remarks: Divisions amongst the various genera of 
conoeoryphids tend to be ill-defined 4ue to 
description ol cranidia alone or because many 
remains of these thin -shelled it ilobne* ore markedly 
distorted, Furthermore, workers have tended 10 
stress thai the group is polyphylclic (Westergard 
1950. Sdzuy 1961; Hutchinson 1962; Kasctti I9f>7) 
On rhe basis of the evident positioning of ihe facial 
Miture on the brim, the present form is a line' 
conoeotyphid (Figs 2 & 4), 

The family Conocoryphidae is rare in the Early- 
Cambrian and much more common ejobaily in the 
Middle Cambrian (korobov 1973). lochman-Ralfc 
&. Wilson (1958) considered that rhe blind 
Conocoryphidae and Hodiseidae were characteristic 
of extraeraronic, enxinic bio fades in (he Cambrian 
uT eastern North America and suggested that Ihe 
blind condition was, favoured by some important 
ecological factor. 

Genus fvshirttcl/us korobov, 1966 

Remarks; The better preserved of Ihe ordinal 
specimens (SAM P24321) is described as having a 
rearwardfy directed furrow on (he side of rhe 
anterior pari of the glabella (.la^o « a/, I9S'4|- 
medially this pan of the glabella seems io have been 
eniite and undivided. Though the front ol ihe 
glabella of ihe new specimen is crushed, there 
appear to be fnur pairs of glabellar furrows (Fig. 
3); the glabella tapers anteriorly and is somewhat 
constricted at about a third its lenutli. These 
characteristics and other aspect* of the eramdium 
conform to the diagnosis of the Karly Cambrian 
genus Ivstunieilus Korobov, 1966, Most Cambrian 
eonoeoryphid genera show three pairs of glabella 
furrows (in advance of the occipital groove). The 
distance the glabella reaches forwards towards Ihe 
frontal border, the possible presence of a raised pre- 
glabella lobe, and the width of the frontal Held. \i 
it is developed, arc snnie of the significant character- 

istics tmpoitant for distinguishing eonocoryphids. 
While Korobov (1966) compared his new- genu* 
with the Middle Cambrian taxon ttailiella Mathew; isMmilat to ihe Early Cambrian PseUd&opi 
I uke, 1940. Species of Pteutfatops may show four 
pairs of glabella furrows, bui the glabella is onl> 
weakly tapered forwards and differ* from that in 
Ivshinieilus fa encroaching on the frontal bordet, 

r^- ■*_■ ,,,x Mm m 


.• .-- ■ -, . 





HS' }• Keprescntiiiion ol ihe glabella tin J :cnti»il pan itl 
I ho I'i'rtnidnini o| SAM P24X-U sltouiue pChilttUl of > he 
four pans til ' ylabeJUu Uitiow.s, 1-4. in advance of ihe 
occipital furrow, occ The irtoie anterior gluheh.u lobo* 
Ml %irunsjt\ eompre«sed in an anitriior^posteiini 
ilirecnoi>, c amp** i\kIi r,L,iie 21$ 

hshitik'tius hriniufuihi sp. \\0\ ! . 
FIGS 2-4 

Conocoryphid sp, indei. I.ieo W at, 1984. pp. in>i-2ll> ( 
figs, 2n-t\ 

Ety/riotoxy: Honouring rhe late LH Brian Daily 

11931-1986) for his contribution lo Australian 

geology and leaching. 

Dtannosts: Brirn weakly inflexed medially, erani- 

dium encircled by border furrow and rather evenly 

covered by fine to coarse, pointed granules (bpm- 

ules) in immature specimens, glabella almost 

smooth in more mature examples; pleura with 

aeieulai, weakly dcflexed terminal spines; eleventh 

sc^mcnJ maciopleurotJs. 

Maivriuf; Holotype SAM P24321, lucking rearmost 

parts (Jaeo $t al. 1984, Uy 2n, b); two paiatypes, 

SAM 1*24322, rear thoracic Moments and 

fragmeniiiry pygidiciru (Jago er at, 19JU fii'. \); 

SAM P24844, eramdium and parts of 14 thoracic 


fM'MTifaittn. ft is unnecessary io repeat the 

descriptions ol ihe holotype and second (Yajemeni 


H I 1- H'WfcilNS/k P. HASLNOHR 



h.>. 4. Keeoivmicfion -J*' L'C|ihtflon tilivJ nun pari cm 
"iiuMMs iii /i ^huiH'Hus brlundatfyi sp. nut , a bom -4 
limes Jimcn v -»."'A 0' bolorypo. 

provided in Jago Of a/. (1984, pp. 209-210). 1 he new 
specimen conftrnis the arcuate shape ol the 
ccpfwlon hefurt\ and slight ntpfcUafl inflexion of ihc 
border, as well ft9 indicating that the facial suture 
h confined to the brim, Jn the holotype, ihe 
granulose brim is poorly preserved and is wider than 
might seem obvious in tbr 4 previously published 
figures; its posterior is well defined and che wide 
border furrow more finely and spaiselv granulose 
lhaii llie central parts of the eranidium. I he new 
remains slum a neatly smooth, narrow glabella 
tapering forwards, wiih lour pairs of glabella 
I arrows directed rearwards. The fragmentary base 
oTctn occipital spine is present in the holutype. 

The additional individual indicates at least 14 
IhOiaelc segments and confirms thai the eleventh 
is inacroplcutous. The thin pleural ridges nn the 
holotypr show a line of 10— J I spinulesor pointed 
granules on the anterior segments and 8-9 such 
gnitiules nn die posterior ones; other fine granules- 
occur beiweeu. 

MvusHtVwettnr: The hololype, SAM P2434J, 
indicates an animal exceeding 25 mm in length. The 
distorted eranuhurn itf SAM 1*24844 is 50 mm wide 
and the maximum widih of ihe glabella is 8 5 mm; 
louiih luotacie segment uboul IM9 mm wide. 

excluding the spines and wiih ihe axis about H).2\ 
the width of the segment; animal in excess of 55 
mm long, 

Comparison: The genus includes two previously 
described forms, Ivshintellus nikolit and Ivshink'llus 
putulus Korobov, 196b, both from the Aldanian 
Stage in Tuva, southeastern Siberia, within the E/im 
Suite on the Ezirn River. Both are represented by 
distorted cranidia and so any comparison with the 
present distorted remains must necessarily lacX 
precision. /. hriundailyi resembles /. ntkoltt in the 
-nape of the cephalon. save for the distmciive 
medial inflexion ol the frontal border; the glabella 
also seems narrower. /. pafafus differs from both 
pf the former in Hs pointed glabella and the 
somewhat quadrate shape of the fixed cheeks. 

Mr J. (j. Gelding has kindly provided 
photographs of cranidia and loaned several nearly 
complete dorsal exoskcletons of eonocoryphid 
trilobites collected from ihe Parara Limestone at 
Bunyeroo Gorge in the Flinders Ranges. This 
material is of a laxon different to /. briandaiiyi as 
the eighteenth thoracic segment is macropleutous 
not the eleventh, and the medial thoracic segments 
have remarkably long and srrong pleural spines. The 
lather long glabella, which reaches a depressed 
border furrow behind the medially widened border, 
has three pairs of incised, baekwardly directed 
furrows and a weak pair of depressions on the 
extreme anterior poriion; laterally, lUe facial sutures 
cut off the major part of the brim. In its ecphalic 
characieristics this form from the Parara Limestone 
shows similarities to the Early Cambrian genus 
Ataps Lmmons, 1844. 

Conocoryphid gen. el sp. indet. 

tiemorks: The additional fragmentary ihorax (SAM 
P24845) does not show ihe eleventh segment 
(counted from che most anterior segment preserved) 
maciopleurous, and the pleural spines are more 
deflexed and evidently shoriei than in the preceding 
form. However, the pleura show the characteristic 
wide-flat pleural grooves and narrow pleural ridges 
of conocoryphids. The thoracic segments are about 
11.3 times as wide as long, excluding the spines, with 
the axis about 0.24 the width of the segment. The 
poor preset vation has largely obliterated any surface 

Disarticulated thoracic segments associated with 
cranidia ol Cienoce/fhtus Hawlc & Corda, 1947, in 
material from the Middle Cambrian of France* 
show the pleural terminations narrowed abruptly 
and thence produced into a somewhat deflexed, 
slender spine. The shape ol ihe pleural terminations 
in SAM 1*24845 is markedly similar. This specimen 
could be .i dimotphie individual of /, briahdaUy* 



or even represent a separate conocoryphid taxon, 
but basically is too fragmentary lor determination. 

Age of irilobitc finds 

Trie occurrence of Iwshinicllus on the Fleurieu 
Peninsula is of potential significance with respect 
10 the timing of a major straioieetonic development 
within the Adelaide Fold Belt. Cambrian rocks of 
the Normauville DfOUfl below the Heathcrdalc 
Shale arc of both basinal and shelf aspect (Daily 
19B3< 1976; Jago et at. 1986), The trtlobites occur 
in the upper member of the Heatherdale Shale high 
in the section of the formation exposed (3aga » j - af 
l£84l At Cafrickalinga and more southerly parts 
of the Fleurieu Peninsula the Healherdale Shale is 
abruptly succeeded by up to ca. 1300m of turbtdkes 
of the Cainckalinga Head Formation ( lata 
of Daily & Milnes 1971, 1972; Milnes 19&H the 
basal interval of the thick psammitic and/or politic 
sediments Ol the Kaninantoo Complex. Sandstones 
of subttdal or lidal origin (Haekslatrs Passage 
Formation) succeeding the Carnekaltnga Head 
Formal ion record a phuse of basin filling. Jenkins 
(ls»K l >| identifies t\u< overall basinal deepening qq<| 
shallowing as the *lvamiiidjeii/.Jadliauiastiaiuiecl- 
onie cycle" and I he organic-rich sediments (Fosk-r 
et ai 19X5) of rhc Heatherdale Shale may be 
considered aa the Vuslule" phase marking the start 
of rapid .subsidence associated with renewed luho- 
spheric extension, 

The likej) agcal hshuiultut In Siberia is teu 

Aldarnan -, :r\ Jagocftf/. 1984) Dl mpili,il 

Early Cambrian, and the comparable eenus 
/',» ,,,.. iin it-; ?*nid Fatly Cambrian 

01 •:.<■ ica and the British Isle* The 

present fin.t.Tu- ihal / h r tj n r/.Ji / Vl and a 
eonoeofyphid frata Buuyeroo Gar$e arc dlffemri weakens tJw ckmu of Jajro et ai (1986) that 
loihv.wvpitid.-. uiggeM a correlation between the 
Hcflthcrdnk Sbik ! I Iwr part* flf th< P*wti I 
i nu'.Mi'Mi .n tno Hinders Ranges, ihoush on a 
testa i ,: ,■ . -»i,sci ..I .i. ... ■ i dark, 

nodular offshore farics ol 1 he loiicr loi malum may 
wcN haw ranrasponsfed wilft Ibe deepening sijmaUed 
hv the HeanflmflaJe Shale he* Jenkins' 1989) 

Trace Rttftfi 

An aupi | . :, | » »l -bice 

Ut.vci- well preserved irirm.pnd i, 
< > a dtclamv of 80 em, parK ot '-v 
Othci tia.i' pruhublj made by bug? uQobi ■ 
a poo;!-, preserved section oi t>% . mm 

prodUwfl bv a florin lily* creature n , 
surface hav been preserved due lo an innuml 

The arthropods evidently moved over a relatively 
fine-grained and soft, muddy substrate with the 
ends of their limbs making deep impressions. Had 
this surface been subsequently buried by further sili. 
the cleavage now present in the rock due to 
deformation (Cambro-Ordovician Delamerian 
Orogeny) would probably preclude splitting along 
the particulai bedding surface. However, the traces 
were buried by a 3.5 cm thick bed now compi ising 
intergrown phyllosilicates and minor fine (-0.1 rnm 
diameter), angular quart/ grains, which are 
commonly amalgamate and show undulose 
extinction, The quattz is probably dctntal. XRD 
analyses of the rock confirm visual observation that 
the bed is in homogeneous with a (less weathered?) 
fraction comprising predominandy illite JM.i 
minor muscovite M {> and kaolinite-smectitc, and 
a second (weathered ) fraction including illite and 
Ca. montinoiillonite. Iron rich opaques are 
concentrated in parts. Illite 2M is consistent with 
a pangenesis involving deep burial and alteration 
of possible older mineral phases. The bed is 
continuous along Write OWE an exposure distance 
Of 60 'ii and thickens slightly towards the south (up 
to 4.5 cm), in places, obvious granular components 
that are apparently 'relicts' of the original 
sedimentary particles grade in si/e from » 0.5 - 
13 mm near the base of the bed 10 - 0.1 mm ai 

thr. | op 

Tta granutai Aggregates of phyllosilicates 
pof&ibl? repraenl the highly altered components 
nf .in air- fall tuff thai mixed with a .small amount 
of deiriial material and formed a slurry which 
buried and moulded the irflobttfi iraecs. During 
modern feathering a htu of parting developed 
SUbjaceni lo rht 'jnileriyingsiiNonesueh that a very 
thin veneer of granulai nifltei ial slill adheres to and 
eompictcty etivei n i|« ita 

The tiail traLrd .:vv Vr SO Cm consists ol SUb 
parallel rov,^ oi rnthef closely spaced imprims Of 

ambulaiory limbs and averages 7^ to BO mm in 

*idiii The ind ( \idtial imprints tend ro be sei 
obliLjiu-lv or L*ncebeIon and the best preserved arc 
about IfJ mm long, I /us uses ^ o\ (h« forni 
illustrated by Seiiucher ti'/5s, fi^, la, b) and was 
evidently made by a peramtoitlafu^ nilohitc with 

Irs b.idv BiXlS alined in the diwnon of fiavel. In 

i »rei ii is svaiapped by a crwdcly scgmt-nicd 

"iM- I'V B ywm Nice atumal. Nearby 

exj^osare^ re the HealhrrdaJe Shale show much 

beD * i sampfaoi probaW>Mhi9 

K tBltOtl v.ilhnunifroii, 
buuows nl a tt.a-iwly brge Inrm of Plonofites 
Nicholson 1)01 i i,nlcd trace !•- pn.hably 

ol endogentc .-u preserved -vampies 

show a rt^eular back-filling of larpe ovate faecal 
mjeed liansvetsclv 



• v - 

I ijt, Bedding plain- below full horizon, showing IrHOC lossik made b> Inlnbiles, A-C, ami sImkIimc, I). atluluiU'd 
loan unknown uoi in-like animal. A-A', trace made by perambulating trilobiic with bodv aliened in direction 
DJ navel; B-B', marks made by large* irilobiie moving obliquely; C'-C, indication ol a tritobitc I ravelling tvl.niwlv 
quickly. Scale bar equivalent Jo 20 cm. 

** "5c * A *** 

Fits. 6. Repivse.uaiioi! of bedding surluee portrayed in Tig- 5 indiealing (hv freVcMl sas 0( marking- made by irilobnes 
]\-\ t \\-\\ , t -t J and another unknown animal (D). 



The two other (races show reta(ivel> widels 
spaced series of imprints interpreted as the 
impte.vaons of iftC tips of the tclopoditcs of 

trilobitcx. In the larger Lrace. the individual imprints 
aie up to 20 mm lont> and on one aide of I he 
complete nail ait arranged m rather h tegular rows 
posilioned in an emeehelon manner. 1 hiv sort of 
maikum is well explained by Setlacher (1955, He. 
le. )') and indicates a lulobilc iikaiiiu obliquely 
crab-wise, perhaps partly in response 10 a ventlc side 
cuticut. I he tips of the limbs which formed the 
marks on either vide ol the naee spanned 
ii|iptox*noatcly 2^ em, indicating ;• Quiltf tyrgt 
aniniat. i he imprints of the remaining tnivc arc 
widely spaced and suggest a smaller am ma I 
travelling relatively quickly with the body uhgncU 
iu tiie diteetum of travel. Jhese marks are not 
especially distinctive {and were onlv seen in certain 
lighting I bui arc comparable \s ilh some ol the lame 
lilooileltailsocc lining in Hie late EatU lo Middle 
Cambrian knngaioo Island (Jioupof DaiU (1956). 
on lie noirh coast of Kanyamo Jsland. 

(he body fossil remains of the trilobitcs described 
liLt'.'in OCClirrcd respectively 1.6 m siratiyraphJcally 
Mow (he surluce with trilobite trails (SAM l*24K44) 
and KK cm above this sm lace ISAM P24845). The 
naccs surest thai Irilobites weie live inhabitants 
ol the hoitoin environment 

Palwobiological signifiennce 

The thmly laminated arjiillites of the upper 
inemhei ( - 240 mm thick) of the Hcalhmlalc Shale 
are almost etiiiielv flat bedded and certainly 
accumulated at depths below storm wave base (cf. 
Jauo et of P)Kr>). A single r ippled surface observed 
may tellect localised bottom currents. Consideiing 
i lie ihtekness ol the succeeding fuibiditcs, the depth 
of water in the depositionni environment was 
probably of lite order of hundreds of rueircv 

);wo cf at. (1984) cite and illustrate apparent 
mudcTack.s in the Heatherdale, Shale liheii fig. 4l 
as possibly indicating a shallow environment o£ 
deposition bul ihev concede that the cracks mas 
rmw formeLt due lo syuacicsts. DJufyhski & 
Walton ll%5l illustraie l*hcir ligs 112 &. 113) 
computable cracks in turbidites.. 

Trihtbtle iiuilsoceur abundantly in parts of ihc 
late Early le Middle Cambrian sequence or. the 
north coasr of Kangaroo Island (Spris.*^ 1955; Daily 
ft a/ 197*?). Examples ol such irails up lo IU— 12 cm 
in width are especially mimeinu.s in intervals of 
these rocks deposited in tidal and subudal faciei. 
Thi* association ol tritoPitc nails in shallow -water 
sediment* is common world wide during the bitter 
l*alaenAue mid eonstiralcs the CnKntmi Facte* of 
Sedachei (I9fvl). I'tluitl is apparent that trilobitcs 

had an ordinary oxidative kind of metabolism such 
a< is elVdTacierisiic of modern, free livinc arthropods 
and mctazoans in general, 

The aithtupod irails m the Healherdale Shale 
suggest relatively larce animals moving over the 
bottom in some numbers. I he intplicaiiou is that 
lite bod> fossil remains o\' trilobiles were part til 
an indigenous population and not isolated 
individuals somehow transported into thcenviion- 
nient by chance. PrCMinmiL! an oxidative meta- 
bolism for (lie (rilobiics, anil the worm-like creatures 
responsible for -.anous burrowed horizons, the 
boltom walet> of lite Healherdale Shale 
environment were clearly not uniformly anoxic and 
most have been at least moderately well oxygenated 
&t the tunes of btoturbaliou. I his contradicts the 
commonly held opiniun thai black shales 
characterise oxygen starved basirn; fluctuating 
oxygen tensions ma> be more likely i;Moriis 1980; 
Kauffman 1982; Kikpcrt 1982; Wet/el WjSH 
Conway Morris I98H 

By meiisunng ihc displacement ol a eonvenicnilv 
sized plaster model oi a irilobile, the soft tissue 
mavs of the animal which made rhc 23 cm wide trail 
may he climated al - 500 9, With reletence to Rp. 
8,(t oi' Alexander (1979), the o\ypen consumption 
of an animal of this size may have been about 0.015 
ml gin lir . or for the whole organism, 23 ml 
hr . Runneiiar (1982) demonstrated then the 
sui foeeaiea oi tlieiespiialoiy organs (e.^;. yills)of 
aqueous animals is approsimalcly propoiuonal lo 
the body mass; this suggests a respiratory exchange 
surface of - 500 cm for the trilobite. Application 
of equation l.\5 of Alexander (1979. p. 2791 
indicates that the partial pressure between the sea 
water and blood over the respiratory surface needed 
to supply the tequiicd osv^en consumption is -0.06 
atm. AsMtminy thai (he respiratory piement in the 
animal's blood was saturated at 0.02 atm, the 
minimum oxygen concentration in lite sea water 
neecvsary for the trilobite was - O.0K aim or - _Wo 
PA1 ( -UQ mm Hg of ,). This \alite lies within 
the medial pan of the spectrum of oxygen conceii- 
naiions m the oxygen nnuunmu layer v\ tniHlcm 
oceans etfeciivelv 2,5'Vn | f AI. to 6(V»'o PAL (Barnis 
& Huchev 1982/fir 1.9), 

There is cunvmlv a renewed wave of speculation 
concfrniu^ the possible increase of aimosphcric 
oxycen do'inp, the laic Prceamhnau and its 
relationship lo t lie emergence ot the meta/oa and 
higher plant lite (Runne^ar l9S2a, b and pcrs 
comm.; Glaessner 1083, 1984), The present example 
sujigesl that simple ohser\atum ol dark coloration 
in pans oi' Precambrian sequences (c.>i ( arcy and 
bhick phyllilcs in the " ? T(JlfrCl8*4ft0 1 Scries 01 'he 
Adelajdcail) is unlikely to aive reliable tnlormatiiM: 
mi ambient nxywen it-nsions. The coloration of 



(halCS will also reflect diagenetie processes tad the 
effects of incipient meiamorphiMn; lor example 
leerystallizahon of" haematite to spcctilarite may 
cause beds ro become almost black (Clemmey & 
Rftdharu I982J- 

/. briifttditifyi k remarkable for the long gena! 
-spines, the attenuated .spines on the pleurites and 
its macropleirrous eleventh segment Similar lone 
spines in ndontopletirid tnlobites have been 
considered to have functioned in helping to support 
the animals on ihe sea boliom, either durinje 
"resting* or while I hey were feeding with the aid oi" 
currents passing between the substrate and the lowci 
side of the thorax ( Wittin^ion 1956a, b; Clark son 
1969; for a different opinion see Rergstrom 1^73 j. 
Such, an adaptation h consistent with /. bnandmlyi 
inhabiting a soft muddy bottom. JJetause of the 

lack of srreamhmng brought about by the spines 
the trilobiie wa\ probably not an a*de swimmer, and 
tins accords with the finding of trackways in close 
proximity to lis remains. However, the track wavs 
indicate larger animals than the bod> in^il 
specimens and there is no way nl telling whether 
the truces were made by more mature individual 
of eouocoryphid.s or other uilobitcs known ui teach 
large size locally, such as Hedhvhiu Cossmann. 1902 
fsee Conway Morris & Jenkins 1985). A host ol 
modern day creatures have adapted io inhabn naves 
and other dark places and become secondarily 
blind, and loss of sight in coiioeoryphids would 
have posed no disadvantage if these prima! denizens. 
of the deep were bottom dwellers who lived below 
the effective penetration nl light, sustained 
primarily on settled derriius 


Au.nanoik, R. McNmi i . (P>79) "The Invertebrates'' 
(C'amhndac ttniv^rsny Press, Cambridge.) 

HAKKrs, R. S, K Si ITUOIUIK, K. N. <l'>* 7 ) "An 
lutpotluctlon »a Marine rkology" (Hlackwelt, Oxford, 

fiuRrtKi RflVT, t |IQR3J Oiganuuuon, life and v*sienMitc-.s 
t>4 HilMhiiCj. Fossils aitd Strata 2, t— 69 

CUkkmjs. h. N. K. 0969) A ttinciionaf siudv fll ihe 
Siliirum odontopleurid oilobhc leonuspis deOexu 
(l.;Ue) tctMa I, J29-.H*. 

< ir-MMh, H. & lUbiiMi. N. |I4.X2j Oyyjjcn in Ihe 
Precamhrian atmosphere: an esalualion of the 
geological evident, Geotogv 10, I41-J46, 

<:unwav MoftftlS, S. (I985J Cambrian lagerstatten- their 
distribution anil sinnificanec. PfiU Vans, /?. Sot Umd. 
B3II, 4s>-fi5 

Si JfsiKtNS. R .1 K (1985) Healed injnucx n ( 

Ftfrlv Cambrian trilohites tin in .Sotilh Australia. 
Akherinza % Ih7-I77 

D\ns, a. mShj 1 lie Cumbrian in Souih Australia, hi T.| 
si>teina Cambrico, su palneogMfm ye el 
iu ba:-e" 20ih J fit. gcoi fctf/ffir, Mexico 2, 91-142. 

liyfO) I he rovjilit'eioiif. Cambrian succession on 

Mriiritii Pemm.itta. South Australia fee V 4M7, Mm 
K 579-601. 

»I976) I he Cambrian ol the I linden Ranges. In 

H P rtKniKon, Bl, Daily, R. P. Coais. A b. P., Forbes-, 
"I,;iie Pteeambrian and geology of the 
.Adelaide Treosvncline" ^ind Siuarf Shell'. Souih 
AuMrnha". i^rt hit. f*eoii Cbnfir. Hxcumion Giade^o 
>?A. 1-56. 

AMjinis, A. R. (1971) .Sliangraphic notes on 
Lower Cambrian t'ossiliteious between 
Oviipbell Creek tind TinkatilUi Death in ihe type section 
of the Kunmantoo (".rrjup, rteurieu f^ntnsiila, Soulli 
Aii5lr;ilia. Trans. H. Sac. S ,\n\l V5, 109-214. 

&. _ 11972) Rovix«on of die slral (graphic 

niimenclaiuif or ihe Cambrian Kdiimanioo (ironp, 
South Ausiratia. J. Geot. Sac. Avit, !■>. 1^7-202. 

^ .TwiiHtr, C K St. BGpkNt, I. A (1979) 

1; Geology mid ecoinorphi)logv. pp I-3X hi M I Uter, 
C R.lU'idale& J. K 1 .ina, tFds) "NaiumI HiMory <it 
Kangaroo Island" [Rpyfl] Sociciv of South Auvuaha, 
Adelaide ) 

IX : i-i-i^M, S., <M Wai i" . I . K, ||9fo) ".Sevlimeril jrv 
t-e.inire^ or Plysch and Ui'-vw.iaiV Di'seloptnenis in 
Sdhmcniotoct 7 (Hsevier Ns»llurlflmls,1 

Vo$iti<, C. B.. C-Tknovskis, A.. & O'Bkii H G. VV. |J99$) 
Oruanic-walteU niierotbssiK from the KrtiK Cwzibriqo 
or South Australia. Alvhermgu «, 259-268. 

Cri ^ESSver. M. I. flWO) The emergcrwe of Mei^Ofi In 
the early histoiy oMTft. Precoma R& 20. 427-441. 
II9S4) 'Trie I (K\ A Biohisumeal 
Stiidy'\ (Cambridge Lniversiiy Press. Cambridye.i 

Hi'Taii.v,</N. K. I). (1962) Cambrian srraiic.uplty ...kI 
trilobne faunas ol southeastern Newiau.idU.nd. Suit. 
Gaol Stj/v Can 88, 1-156 

J.soo. j. B.. Daiiy, a, VitN Dck DckLii. C C () 
O.KS<.iv>KN, A, A Sal-M>h<s, N. (|^H4) lirvr reported 
irilnbfres trout the Losver t "ambnan Noi manvilfc Ciniii|\ 
t-li/uiicu Peninsula. Soiiili Ansiralia. Trans, ft. btw S 
Attst. 108, 207-211 

., ot (hi ino. .). <;., a.- Da(i v, b n i )R6) CiWjtn'tin 

scdimenisof the Selltck Hitt-CinvktihTiga Head arra, 
Flcufieti Peninsula, Snutli Auslulut. (ft A. I, Paik»'r 
(Ed.) <l One Day ( "ieologtcal Excursion id ihu \<lelatde 
Kc^ion" pp 67-Sl. rCieolo^ical Socieis at* Auxiroiia, 
SA^ Div, Adelaide.) 

Jl.NMMs, K. !. y (1989; The Adelaide Kilil Belt* tcvl 

teuppraivd. In .1. D. Jago, & K s. Moore, \fi,U\ " Vpec^ 
of the Adelaitlean Slid C'diuhrian fleolpei- \»f ihc 
Adelaide Ijoosynetiiie" (tiul. Sot. Ansl Spev. Pulil 
in press,) 

Km til man, L. G. (iy«2) The eommumrv simuiire of 
'shell islands' on oxyscn deplored frubslfaup in Misozoic 
dark shales anorf laminated curbonaics, Tn Cj. fein*efc& 
A. Seilftcfien (Kdsl 't-'yehe and event siraJificatnorr'tyV 
502-500 (Springer Verla?,, Berlin Hekletbeig J 

Koxobf^, M N <tsi66) New Oilobile< or Ihe inmity 
ConrictirvpiudiK' tiom the Cambrian hf The .Siheiian 
HUiirurm and Tu\a. Pa/ctwt. /h IV6A \o. 4. s>2-97 

— __ . (IV73i The tolobite (amity ConeitorvpliicJac nnd 
its sigmticAnce (o ihe sirai,i)?raphv ot the Cambrian 
deposit*. Trudy .eeo/. Inst, Moscow, 21 1, (In Russian). 

Kisf-cpt. W. (I9H2) r.nvfronmenitd Changes during oil 
ihale depoMiion pis deduced from slabh i>otopc rAiiris 
hi O, tlmsule Ik A -Seilaeher. (Ed.) "Cvclie and eveni 
slrAtdiealion" pp. 482-501. (Spnnger A\rr|au. Bolin, 
Heidelber)? ) 

I ii.'1i,viam-Bai K, C- * \ V "M'%. 1 C, (|958| t.anthrhtn 
biOstr.iiiiiruphy in North America ./. M/rWin/ 33w 



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their relationship to the Cambrian Kanmantoo Group 

pp. 43-52. In A. J. Parker (Eel.) "One Day Geological 

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Geot. Soc, London 137, 157-170. 
Rassltii, F. (1967) Lower and Middle Cambrian trilohiic 

faunas from the Taconic sequence of New York. 

Srtiithsan. tnisc. Collns. 152, l-lll. 
Runmegar, B. (1982a) Oxygen requirements, biology and 

phylogenetic significance of the late Precambrian worm 

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_ . (1982b) The Cambiian explosion: animals or 

fossils' 1 -J. seof. Soc. Aunt. 29, 398-411. 
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Trilobiten. Abh. muth-natunv. K. Akad. Wiss. Mainz 

1961, 597-693. 
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._ (1964) Biogenic sedimentary structures pp, 

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(Eds) il Cyclic and event stratification" (Springer Verlag, 

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Wittington, H. B. (1956a) Type and other species of 

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Odontopleuridae. Bull. Mus. Comp. loot. Harv. 137, 







The first live collected specimens of the South Australian allied cowrie Primovula (P.) heleneae 
Cate, 1973 are recorded. This species, previously known from a single shell collected prior to 1908, 
is redescribed, and details of the animal and its habitat given. Crenavolva cruenta sp. nov., is 
described from Spencer Gulf, S.A.; it most closely resembles C. striatula and C verconis but is 
distinguished by its more elongate form, more produced ends and colour pattern. 
KEY WORDS: Gastropoda, Ovulidae, South Australia, Primovula heleneae, Crenavolva, new 
species, habitat. 

Ifto&ffllorto 01 tb* IfaM SoL'tpy af& Att&t, (1*8$, 11* I) 205-214. 




by K. L Gowmi-HoLMrS* & N. J. C. Hq\ MESl 


OCWJ ii r-HtW Ml i t K IlV Hoi mi s, N. J. C. . (IW» Rediscovery of t'rxmovulu {PJ fh'Unc.ic i .or fthd 
dcscupliuti ol rt how species of i'rfmi\<ilvn born Soutli Ansiiult.i (Mollusex ( Invropocfa: Ovaidae). Twm*. 
A'. .Sue. ,S. Aw.t, 113(4). 205-214. 30 November. IVN'> 

i in* tii ,i iiu* i niiecied specimens of the South Australian allied cowrli Pffotvvufo (ft) hebenea* <ni<\ 

1^71 art nrnKJcd Dns species, prcvloUftfy known I mm ;i single raVII collected pikv to 19Q8* b icdcscribed, 
and details of ihc animal and lis habitat glvcrt, DtfrttfVtfAtf <n/fniu ,p Rot , if di-.a ibrd From Spencer 
GUlf, S.A.; ii mo*l closely resembles C, str'uttukt and C vvrvi'tiis bin is Llbfin|t)i)licd In h rcdoffgjirc 

loiui, nunc produced ends and colour pnitern. 

ki v Wouiv-,; CaMtopoda, Oyulidac. South Ausiraltn. PrtttlOVkfa AtffeftW?, ( 'WWVft/M new sprues, habitat. 


I luce SpCClC5 ol' the allied cowrie family Ovulidae 

aie recorded hom south Australia! Cwiw/va 

ven-anis (Colton & Godlicy, l952>. Pnmovula (P.) 
/;t7tYRwCak\ IS73, and an ricfyvitlva sp. (Coleman 
WKI). All are regarded aS veiy rare. The first two 
species are known from single specimens dredged 
dead pi tor [0 1908 (VCfCO 1908, Btf Ovula fornt<i\<t). 
OH Douglas Point i»i northern Spencer Gulf, 
S. Ausl , one oi' ns (NJCH) round a number ol 
allied lwi ies living on fans of the gorgonian corals 
I ii/)/r\uun; sp. and J.chmoyotxiu sp. in 10-15 m 
depth. These specimens were identified as 
Pnmovula (P) he/erteae Cute, 1973, (living on 
f H/'/c.witttf sp), and 8tl undesenbed species o( 
Crerttnoh'a. (living on Echinogargfa sp-h 1 lere we 
deu ribe ihe new species ot Crenuvolvtu redescribe 
>f rtty heleneae, and include details Of 'he animal 
and its habitat. 

Materials and Methods 

The material repotted here is deposited in the 
South Australian Museum, Adelaide (SAM), 
Australian Museum, Sydney (AM), Museum ol 
Victoria, Melbourne (NMV), Northern Territory 
Museum of Arts & Sciences, Darwin (NTM), 
Queensland Museum, Brisbane (QM), Western 
Australian Museum, Perth (WAM), Tasniauiati 
Museum and Art Gallery, Hobart (TM) and 
National Museum oi New Zealand, Wellington 
(NMN/). living material was collected by 
m. I HVatuI il Hookah"divinuand usingaSmith- 
Mcltnyie grab dredge. Underwater photographs of 
the living animals were taken using Nikonnos 111 
and Nikonnos V cameras with macro extension 

k South Australian Museum. NJortb lerracc, Adelaide. 
Y Ausl. MMH) 

KfllMll Maiine Seienoest 2Afl Last retrace. Adelaide, 
s. x.i.i 5000. 

tubes and electronic flash units Wet material was 
preserved in 75^o methylated ethunol solution CM 
2% formahn/propylerie glycol solution, Uadulji 
were prepared lot examination nudei lean rtf 
electron microscope (SF.M) alter ihe ruetbod Q\ 
Bande] (NK4). Colo it descriptions follow Kornerup 

c<: Wanschci (1*78). 

Primovuia tPrinovula) Iwkt'i < 197^ 

FIGS l. 3, 4, 

Ovttki funitosu Vik >, l9(tH, p. u: {»\ou Adams & KeClfi 

Prostmkta vtxeoni:;: «. otton & Godfrej |412j p (A, i«l 
l. ii- l^ on pact) 

IWtusirnnki ven-nws Irritate, IW, p [OS; COI I^V>, 

p. 368, (in par i ). 

Primovtifa (Prtmmula) wrvonft: Setulder, 1941. g, I0"\ 

(tn part). 

\ ; i'<>':inmia (Pi'//usi/nniuj vcrawis: AJIan. J'^56. p L3t) 

pi, 14, lig. 40 (in part). 

Ptimtnul,, (Pnnynnho hr/rru-ur ( jlc. 197.1.*. p 41-4, 

fe B9, 

/i/v Hotolype (SAM DI5941), part oi ihe i 
series examined by Cotton i<: Ciodfrey (1932); a 
puiutvpe of Prosimnia vercotus. 

type locality: Hotoiype dredged In QuIfSt vntccni 

or Spencet Ciull', S. Ausl., by J. C. Vereo; the 
holotype of P, verronis was dredged off St T rands 
Island, S. Aust.. by J. C. Vereo. 

Other Material: SAM D17973 (4 * e^gs, in spirit*. 

SAM DI7975 (2, dry) Douglas Point, noitheru 
Spencei Gull, S. Aust., N. .1. C, Holmes,; 
SAM DT84M (13, dry), SAM DI8435 (8. m spirit. 
Douglas Point, northern Spencer Gull', S. Aust., 
N. .1. C. Holmes, $m198K 

Species description: Small allied cowrie, aclull length 
9.15-13. 80 mm. mean 10.77 mm t J.(XI4. i-longaie; 
both cuds slightly produced, posleriorcnd moie so; 
widest pari of dorsum in posterior ' ■■'. (Tie }[ ) 



Fig. |. Primovuia (P) heleneae, two adults, one with partially extended mantle, on the 

gorgonian coral Euplexaura sp. 

Fig. 2. Crenavolva cruenta sp. nov., two adults with partially extended mantles on the 

gorgonian coral Echinogorgia sp. 



Dorsum (Fi£>. 3A) sculptured with numerous 
mans verse wavy stride, fuiiieulum prominent, with 
three weak transverse ridges, Base (Fig, 3B) concave, 
outer lip and paiielufcolumeilar lip rial and inward 
sloping, apfcrtflfe narrow with o strong fossula, both 
sides stronyly calloused Patieial-cotumeb'ar base 
smooth but wuh bodv whorl striae visible beneath 
the callus. Outer lip with weak transverse ridges 
posteriorly, inner margin slightly dentate 
posteriorly; outer margin with three to ten slight 
tQ piominent nodules at posterior end-, morc- 
ptominenl in smaller shells, usually with one to 
three slight nodules at anterior end. 

Posterior canal somewhat produced, with a 
basal ly projecting terminal knob dividing canal into 
distinct left and right channels (Tigs 3C, 3D). 
Anterior canal shallow *0J "-shaped, slightly llanng 
and turned to the right, bordered on both sides by 
d iiuct ana ftllgfttty projecting terminal ridces. 

Shell pastel red to pale Orange lb greyish yellow, 
anterior and posicuor ops usually orange; paler 
baud across widest pan ol doisum. columellar 
tallu.'-. OUt« lip and labial callus all paler than 

Animal rnanlle lobes (Fig. U with do papillae; 
translucent pastel red to dark orange, with irregular. 
teddtsh brown patch-, trying in size, area 
immediately around each patch not patterned, 
remainder ol mantle patterned whh slightly raised, 
while dots, Toot pinkish white; anterior dorsal foot 
with a senes ol prominent., vertical lamellae; 
posicuor dotsal font with a longitudinal, vfofel 
tr.-AM to brown uupe. leutaele.. wilt, white op lor 
1/6 kii.crh. mo!-;; brown for almost half length, 
remainder pinkish white with basal 1/5 from 
swollen, with violet brown stripe from eye to base. 
Siphon and proboscis violet brown to brown. 

Radnla (Fig. 3f) with central teeth narrow at 
base; wiiii large, broad, Hal heads, outer edge wirh 
man) perpendicular, sharp cusps, central cusp very 
large, natrow at base, widening Tor about halt 
length, where there is usually a lateral denticle an 
;u least one -vide, then rrnmwlflg aptealb, to sharp 
point! wrlh eight 01 nine small cusps on either side 
q| cctUral one first lateral teeth elongate, strongly 
curved, heads tttcuspidatc, cusps uneven, central 
cusp largest. Second lateral teeth elongate, with 
broad. Hat heads, heads with about live long, 
FUrVftl, sharp-tipped cusps. 

EjiK mass (fig, 4A) with 57 colourless, 
transparent, jelly -like capsules laid on a single 
bram.h of the host sequentially in an cloneated 
spiral and iuscd together fach capsule contains 
many white-shelled embryos, shells wider than long, 
shell lettgSfl I5'*-230^m. shell round, punctate, with 
(tun operculum (lies -JR. 4C). 

Range: The new material examined here is from 
nothern Spencer Golf. S. Aust. Cale ()97Ja) gave 
the type locality Of ft (P.) hcierteae as the "S end 
of St Vineent Gull, South Australia (34°55'S 
I3«S°I0'H)", but (he holotype is labelled in Vcrco\ 
handwriting "G. St Vincent or Spencer Gulf. 
dredoed", so Gate's (1973a) restriction ol the type 
locality i.s suspect. The ran^e o| (his species i, 
assumed to be central S. AusL It is notewonhy that 
Cotton (1935) relets to the occurrence nT 
Cretwvo/i'a verconis in southern W.A., but we ear. 
find no specimens to substantiate it Gate 1 1973 w 
icfers Cottons ilW) reicrenee to a Queensland 
species, PrtmovNfo (Pjphnysia Gate. 1973 Us Qvuk 
forthosa Vcrco), but does not mention if lie 
examined the specimens, 

Hahitoi; Ow tans ol the gorgoman coral Luplexaurv 
sp, (Ctudana: Octocorallia: Plexaundae), in areas 
ol strong tidal movemnu 

Remarks: The specific identity of (he host coral 
(SAM m\\i could not be determined (K Aldei- 
filadc pers. L'umm,), The white spicules i.T Ig. 5C) are 
tvpical of huplpwura. tn northern Spencer Gulf, 
colonies (fi£ 5A) arc mulii branched tans in one 
plane perpendicular to the strong tidal currents, Uicy 
•are attached by a thickened base to a large shell 
'Moment Crf other piece or pieces of solid J. i 
ID 'he substrate; a poorly sotted, mIiv, shelly sand. 
Colonies are relatively large (ca 30 cm Uxro. and 
widely and irregularly dispersed at 10-20 ni depth. 
Distribution appears to be associated whh tidal 
current velocity and substrate siluoess, lupfcyouw 
sp. appears to be less tolerant of silt rh*n EetliW 
£or\>io sp.. and is less common. Each Euplgi 
sp. colony usually hosts |Wo to five tnUiYidnafc of 
P. (Pj lieleneae. Polyps are usually withdrawal 
during daylight on all of rhe colouv JCgurJlrv ,. 
the pivsence of P (Pj Helena. Orientation ol 
shell is always whh the longitudinal a,\r pat tifel rtj 
the branches of the hosr, with na observed 
prelercnee lor a head up ot head down attnudc. No 
particular position on the host is favoured. 3 tut 
adults and juveniles are present toecihc-t Eggs have 
been observed in late autumn and early winter, laid 
on a single branch of the host and not attended by 
the parent, although adults were present on the host. 
The colour pattern of the extended mantle lobe- 
fUj fu-k-neae is almost identical to o\. g 
Euplcxutira sp, The background colour of thccoial 
is pastel ted, and rhc withdrawn polyp's are red to 
reddish brown. The pattern ol reddish brown 
patches on the mantle lobes of P fPj httencac 
mimics the posaion of the polyps on the coral, and 
an animal with the mantle lobe* fully extended N. 



rig. V Priniovulu (P.) helemvc (SAM DI7y75), A. dorsal view, \ 7.5; B. ventral view. \ 7.5; C & D. lateral views, 
x 7; E. size range, x 2.5 (SAM D1S434); F. radula. scale bar = 10 /tin (SAM DI7973). 

very well camouflaged. The very close mimicry by 
P. (P.) heleneae of its host suggests that it is an 
obligate predator and that its distribution will be 
limited by that of Euptexaura sp. The distances 
between colonies of Euplexaura sp. tlO's o\ metres) 
suggest that adults of P. (P.) heleneae would not 
move between colonies, so dispersal is probably by 
a free-swimming veltger stage oi' short duration. 

Crenavoha cruenta sp. nov. 
FIGS 2, 7. 

Holotype: SAM DI8431, dry adult specimen 
10.45 x 3.60 x 3.00 mm, collected on the gorgonian 
coral Echinogorpja sp. at 15 m depth, 1.6 km E. of 
Douglas Point, northern Spencer Gulf, S. Ausl., 
32°5i'48' , S, 137°49U2"E, by N. J. C. Holmes, 
5. v. 1988. 

Paratypes: Same locality data as holotype. Spirit 
specimens: SAM D18433, 14 juv., 4.65-9.60 mm, 
24 ad., 7.35-11.65 mm. Dry specimens: SAM 
D18432, six juv., 6.30-9.15 mm, 27 ad., 7.15-12.35 
mm. SAM D18437, 11 ad., 6.70-12.60 mm. AM 
C156094, two ad., 9.55 and 10.90 mm respectively. 
NTM PI 384, two ad., 9.45 and 10.20 mm 
respectively. NMV F52836, two ad., 9.35 and 9.95 
mm respectively. QM MO.19385, two ad., 9.40 and 
9.70 mm respectively. TM E17744, two ad., 9.15 and 
9.35 mm respectively. WAM 152/88, two ad., 8.30 
and 10.30 mm respectively. NMNZ MF. 49494, two 
ad., 8.65 and 10.60 mm respectively. 

Other material examined: 1\vo broken specimens 
in spirit from Douglas Bank northwest, northern 
Spencer Gulf, S. Aust., 32°47'18"S, 137°49'12"E, 
E. Oks, Sept. 1987 (SAM D17976); Backy Point 



I fa \ Phmovulu (P.) hrleneae «S,\M DI7973), A. egg mass on branch of Euptexaura sp., x 4; B. embryonic shell 
dorsal vjcw; C embryonic shell, showing apenure and operculum, scale bar - 10 ,<rn. 

cast, northern Spencer Gulf, S. Aust., 32°55WS 
BTOiWft E. Oks, Sept, 1987 (SAM D17977). 

Diagnosis; Small allied cowrie, adult length 
6,70-12.60 mm, mean 9.51 mm ± 1.2. Elongate; 
both ends produced. Dorsum sculptured with 
numerous transverse wavy striae; Funiculum not 
prominent; aperture narrow with strong fossula; 
outer lip outer margin with slight to prominent 
terminal nodules; both canals bordered by slightly 
projecting terminal ridges. Shell greyish orange to 
brownish red; tips red to orange; dorsal bands and 
patches usually present, white, pale yellow to pale 
orange; columcllar callus, outer lip and labial callus 
yellow to li^ht orange; fossula pinkish white. 
Animal mantle lobes translucent pale greyish pink 
with dark red spots and small and large white 
papillae; foot ruby, underside with median 
longitudinal very pale pinkish white stripe; tentacles 
with white tip for W length, remainder of tentacle, 
siphon and proboscis ruby. 

Description of Holotype: (Figs 7A-D) Shell 
elongate; both ends produced; widest part of 
dorsum in posterior 2/5. Dorsum sculptured with 
numerous transverse wavy striae; funiculum not 
prominent, smooth. Base somewhat concave, outer 
lip Hal, inward sloping; parietal-columellar lip 

convex; aperture narrow with a strong fossula, both 
sides strongly calloused. Parietal-columellar base 
smooth with faint transverse striae present on 
central inner part. Outer lip with rounded transverse 
ridges, becoming weaker anteriorly; inner margin 
smooth, outer margin with four slight nodules at 
posterior end, three slight and one prominent 
nodule at anterior end. 

Anterior canal shallow "U "-shaped, slightly 
flaring, bordered on both sides by distinct and 
slightly projecting terminal ridges. Posterior canal 
somewhat produced, shallow k, U "-shaped, strongly 
Flaring, bordered on both sides by distinct and 
slightly projecting terminal ridges, outer lip ridge 
terminating in a nodule, parietal-columellar lip 
ridge with two nodules, one terminal. 

Shell orange red, anterior and posterior tips 
bright red; light yellow median band across widest 
part of dorsum, continuing onto base; light yellow 
diffuse band midway between median band and 
posterior end, continuing onto base; light yellow 
patch almost midway between median band and 
anterior end. Outer lip and labial callus yellow; 
columellar callus translucent yellow with dorsal 
colour pattern visible. Interior of posterior and 
anterior canals red, becoming orange red at very 
tips. Fossula pinkish white. 



fig. 5. A, Euplexauru sp. colony in siHi, \ 0.8; B. Echitiogorgia sp. spicules, scale bar = 100 ^m (SAM H356); C. 
Euplexauru sp. spicules, scale bar = 100 y.m (SAM H355). 

Etymology: "Crural us" (L.) meaning bloody, 
pertaining to The main colour of the shell and the 

Additional Characters from Paratypes: Animal 
mantle (Fig. 2) lobes translucent pale greyish pink 

with random, dark red spots, rows and groups of 
small, simple white papillae and occasional much 
larger, white, elevated, rounded papillae with wart- 
like knobs, mantle more opaque white at base of 
papillae. Foo! uniform ruby, underside of foot with 
median, longitudinal, very pale pinkish white stripe; 



Fig. 6. Echmogwgte *p. colony /a %//u. \ 0.75. 

anterior dorsal foot with scries of prominent, 
vertical lamellae, becoming less distinct posteriorly, 
disappearing about ', i length of fool from anterior 
end. Tentacles with white tip for Va length, remain- 
der ruby with basal '/'& from eye swollen. Siphon 
and proboscis ruby. 

Radula (Fig. 7F) with central teeth narrow at base, 
with large, broad, flat heads, outer edge with many 
perpendicular, sharp cusps, central cusp very large, 
tapering apically to sharp point, with sharp, lateral 
denticle on either side at about half length; with 
five or six small cusps on cither side of central one. 

First lateral teeth elongate, strongly curved* heads 
tricuspidate, cusps uneven, central cusp largest 
Second lateral teeth elongate with broad, flat heads, 
heads with eight long, curved, sharp-lipped cusps 

Variation: Paratypes differ from the holotypc in 
size, shell colour and sculpture (Fig. 7F). Shell 
colour is from greyish orange (o brownish red; 
anterior and posterior lips red to orange; dorsal 
bands and patches white, pale yellow to pale orange, 
and vary from very prominent (often forming three 
distinct bands) to almost absent, median band being 
most conspicuous; columcllar callus, outer lip and 
labial callus yellow to light orange. 

Funiculum vanes from smooth to with up to five 
weak nodules on inner edge. Outer lip outer margin 
nodules vary from very slight with one prominent 
nodule at anterior end, to three prominent nodules 
at posterior and two prominent nodules at anterior 
end. Posterior end usually much moie produced in 
small specimens, almost rostrate. Posterior canal 
parietal-columcllar lip ridge occasionally with three 
or four nodules. 

Range; Northern Spencer Gulf, S. Aust. 

Habitat; On fans of the gorgoman coral 
Echinogorgiu sp, (Cnidaria: Octocorallia: 
Paramuriceidae), in areas of strong tidal movement. 

Comparison with other species: The new species 
belongs in Crenavoha, which is distinguished from 
other genera in the Ovuljdae by the small, elongate 
shell with median angulation, narrow, produced but 
truncate ends, dentate, curved aperture, striate 
dorsum and ridged outer lip (Cate 1974), C cruenta 
most closely resembles C striatula (Sowerby, 1828) 
and C vervonis, the congeners recorded from 
Australia. Cate (1973a) records two subspecies from 
Australia, C s. traiUii (A. Adams, 1855) and C s. 
tinctura (Garrard, 1963), both from Queensland. C 
cruenta can be distinguished from C striatula and 
C. verconis by its narrower, more elongate form, 
more produced ends, the shape of its more elongate 
aperture, and shell colour pattern. 

Remarks: The specific identity of the host 
gorgonian coral (Eehinogorgia: P. Alderslade pcrs. 
comm. SAM H356) could not be determinded; the 
red thorn scale spicules (Fig. 5B) are typical for the 
genus. In northern Spencer Gulf, colonics of this 
species (Fig. 6) are sparsely branched fans in one 
plane, attached by a slightly thickened base lo shells, 
fragments or pebbles in the substrate; a poorly 
sorted, silty, shelly sand. Colonies are relatively 
small (ca 15 cm high), and appear much more 
tolerant of silt than Euplexaura sp., being widely 



Fig. 7. Crenavolva cruenta sp. nov. holotype (SAM D18431), A. dorsal view, x 6.6; B. ventral view, x 6.6; C & D. 
lateral views, x 6.6; E. size and colour pattern range, paratypes, x 2.25 (SAM D18437); F, radula, paratype, scale 
bar = 10 ^m (SAM D18433). 

but patchily distributed in 10-20 m of water, often 
in quite high densities (up to 20 colonies/m 2 ). 
When C. cruenta is present, each Echinogorgia sp. 
colony usually hosts one to three individuals. From 
observations made in daylight, the host's polyps are 
usually withdrawn on all or most of the colony 
when C. cruenta is present. Orientation of the shell 
is always with the longitudinal axis parallel to the 
branches of the host, with no observed preference 
for a head up or head down attitude. No particular 

position on the host is favoured, and adults and 
juveniles are present together. 

Colour pattern and form of the extended mantle 
lobes of C. cruenta are almost identical to 
Echinogorgia sp. The background colour of the 
gorgonian coral is red: the polyps are reddish orange 
when withdrawn, yellow when extended. The thorn 
scale spicules in Echinogorgia sp. protrude through 
the surface of the colony, giving it a rough texture. 
These protruding spicules help retain a ragged layer 

RLD1SCOVFRY nr PUIMUVU! A IP} ft£&fifV£«£ ( AIL- 


of mucus and detritus upon the surface; this 
mucous layer is white, and pafehily masks the red 
background colour. The pattern of reddish brown 
patches and \% hile papillae on The mantle (obes of 
C. crucntu mimics the ragged mucus on the 
gorgonian coral,, and an animal with the mamie 
lobes tally extended is very well camouflaged on 
us hose. The very close mimicry of C. cruenta to 
its host suggests thai it is an obligate predator and 
that its distribution is limited by that of 
tchinoiioriiia sp. The distances between colonies of 
Ecfwiogorgia sp. (often <0.5 m) and the suitability 
of c: (Merita's camouflage suggest that the latter 
would be able to move across the substrate between 
colonies, so dispersal is probably only partly by a 
free-swimming vehger. 


The generic names used in this paper tollow the 
system and modifications proposed by Cate (1973a, 
1973b. 1974). The distinctions between some ol the 
genera used there are inadequate, 

Tfic two species of Ovulidae previously named 
from S, AusL: Crenavolva verconis and Primovitiu 
fP) heleneae, arc both based on damaged, dead 
shells. Vereo (1908) had three specimens and 
recorded them as Ovuia fonnosa A. Adams & 
Reeve, 1848, a species described from Borneo. The 
description and photographs of the holotype of O. 
Juntu>Mt {= Deiortovohv Jomtosu (A. Adams & 
Reeve, 1848)] in Cate (1973a) were compared with 
the holotype of C. verconis, with the hulotype and 
other specimens of P. (P.) heleneae, and with the 
types of C. cruenta and D, formosa was found to 
differ signilieamly from them. D. fonnosa is origin 
purple with yellow tips; the lossula is not prom- 
inent; (he whole shed is much narrower overall, with 
a narrower outer lip than either C. amenta. (\ 
votvonis or P (P.) heleneae, and * more pointed 
puMeiior tip than either C. entente or C. verconis, 
and which lacks the basal !>■ projecting terminal 
knob of P. (P) heleneae 

Conon & Godfrey (1932) described VercoN 

specimens as a new species, Prosirnnia verconis, 

their description obviously was based on all the 

specimens, although the holotype is clearly defined 

as the shell from St Francis Island (SAM D13476, 

= DJ0174). Cine (1973a) re-examined The types of 

P. verconis, which now consisted of two specimens, 

(one specimen of the original scries is missing, 

presumed lost), and discovered that they were not 

eonspeeif'ie. Cate acknowledged the designation by 

Cotton Jt Godfrey (1932) of the specimen registered 

SAM D13476 (= SAM DI0174) (very woni and 

tailed), as the holotype ot Crenavolva verconis 

(Cotton & Godfrey, 1932), and used Cotton & 

Godfrey's (1932) paratype (SAM DI5943), a worn, 

faded and broken specimen, as the holotype of 

Primovnia (P.) heleneae. The live collected maieriaJ 

from upper Spencer Gulf was compared with the 

types of both species and, all hough both types are 

damaged, the material collected from Huplexaura 

sp. was clearly conspeeific with P (Pj heleneae, and 

all of the new material was different from the 

designated holotype of C vercowx. 

According to Coleman (1981), the specimen 
illustrated by Coleman (1975, fie, 262) as Neoshnnia 
verconis (Cotton &. Godfrey, 1932) is actually a 
specimen of an unnamed species ol" Aclyvolva. 


We are grateful to Dr B. R. Wilson for his 
considerable assistance and critical comments, and 
to Mr P. Aklcrslade for identification oi' the 
gorgonian corals, Mr W. Zeidier, Dr C Watts.. Dr 
F. E. Wells and an anonymous referee are i hanked 
for critieal comments. We thank Mr R Thomas, 
Mr D. Evans, Mi M. Moore and Ms J. Young, all 
of Kinhill Marine Sciences, lot assistance in the 
field. The photographs (other thnn Figs 3, 2, 7, 5A, 
6) were taken bv Mrs ,1, Forrest and Ms A. Renfrey 
(SfcM). The colour plates were funded by the 
Research Fund of the South Australian Museum. 




Allan, J, (1956) "Cowry shells of world seas." (Georgian 
House, Melbourne). 

Bandel, K. (1984) The radulae of Caribbean and other 
Mesogastropoda and Neogastropoda. Zoo/. Verhandl. 
Leiden 214, 1-188, pis 1-22. 

CATC, C. N. (1973a) A systematic revision of the recent 
cypraeid family Ovufidae (Mollusca: Gastropoda). 
Veiiger SuppL 15, 1-U6. 

(1973b) The Ovulidae: Replacement names for 

four ovulid homonyms. Veiiger 16(2), 238-239. 

(1974) The Ovulidae: A key to the genera, and 

other pertinent notes (Mollusca: Gastropoda). Ibid. 
16(3), 307-313. 

Coleman, N. (1975) "What shell is that?" (Hamlyn, 

(1981) "What shell is that? 2nd Edition." 

(Lansdowne, Sydney). 

Cotton, R. C. (1935) "Combing the southern seas." 

(Rigby, Adelaide). 
(1959) "South Australian Mollusca: Archaeo- 

gastropoda." ( S. Ausl. Govt Printer, Adelaide). 
& Godkri-y, F. K. (1932) South Australian shells 

(including descriptions of new genera and species). Fart 

111. £ Aust. Nat. 13, 35-86. 
Garrard, T. A. (1963) New species of Mollusca from 

eastern Australia. /. Malac. Soc. Aust. 1(7), 42-46. 
Irldai f, T. (1935) Australian cowries. Aust. Zool. 8(2), 

Kornerup, A. & Wanscmer, J. H. (1978) "Methuen 

Handbook of Colour. 3rd Edition." (Eyre Methuen, 

Schilder, F. A. (1941) Verwandtschaft und Verbreitung 

der Cypraeacca. Archiv Mollusk. 73(2/3), 57-120. 
Vhrco, J. C (1908) Notes on South Australian marine 

Mollusca, with descriptions of new species. Part IX. 

Tram. R. Soc. 5. Aust. 32, 338-361. 


by Margaret Davies * 


The egg mass and tadpole stages of Litoria eucnemis are adapted for a stream-dwelling habit. The 
egg mass is compact and adherent whilst the tadpoles have suctorial mouthparts and shallow fins. 
The eggs have a large yolk reserve used by the tadpole for nutrition between stages 19 and 26 after 
hatching. At 16-24°C, L. eucnemis metamorphoses 74 days after egg laying and 69 days after 

Inwmitom iff the K"V'W Sfuh-ty of S- Aust , (19K l )) ( 1 13(4 1 215-22** 


by Margarht Davils* 


D.-wirs. M. (ly&yt Developmental biology ol the Aiwtralopapuan livlkl t'lo^ Uiono futnftms (Aintr.-i 
HylidaeK Irans. H, Snc S. Ausf, 113(4), 215 -220, 30 November, 1989. 

The egg Olfltt mid tadpole Plages ol / irortu ewrtewis are adapted lor a Mream-dueHinu bahit, I he ecu 
mass is compact and sdhercnl whilst the tadpoles have suctorial mourhpans anil shallow I'ins.. The pggs 
have a large yolk reserve Used by the tadpole tor nutrition between stage l 1 * and 26 after hatching. Ai 
I6-24°C. /, euvhemiK metamorphose* 74 days alter eye. laying and (W day*, rmcr haiehnir. 

Kj 1 Wokiv, : t.itorut cut'nvnus, tadpoles, developmental biology, lotic adaptations 


i.tittriu eitcnetms (I onnherg) is distributed in 
rainforest streams ut widely-separated localities in 
New Guinea (Tyler 1968; Zwcifel 1980) and on the 
Cape York Peninsula in northern Queensland 
(Zweit'el 1980; COggttJ 1986). The species has had 
;iti unstable nomendatural history in Australia 
(Tyler & Waison 1985) and the uam^s IJtorio 
eucfjemis, L nemtta and L tiaumacidatu have 
variously been applied to Ausltalian representatives 
ot I lie species. 

I ifc history data ate confined to a comment that 
ihc c££s are dark brown and dial nothing appears 
distinctive about the tadpole (Menzies 1974). 

A pair of L eucnemis bred in the laboratories 
ot the Zoology Department. University of Adelaide. 
The tadpoles were reared to metamorphosis and the 
development ol the species is reported here. 

Material* and Methuds 

Specimens of /.. eucnemi^ were collected by M. 
Mahony, J. Read, M. Schmidt and T, Haff at 
ChHrmiHin Ck, Qld (I7°4.1 ' 14591 ') in late August 
ls\SS and reached Adelaide on U.X.I98N. A pait ol 
frogs, amplcctant Upon arrival, ssas [ransfetred to 
a jalass aquarium with a pond of water at one end. 
The aquarium wns maintained at room temperature. 

Spawn was removed from the pond, and placed 
io dechloiiualed, aerated lap water ill a glass 
aquarium 25x25x8 cm lor 24hr at 30°C. The 
aquarium was then maintained at toont temp- 
eraiure, aeiuted and the watet changed daily. 
Tadpole* were led on tightly-boiled lettuce leaves. 
The culture was divided ifi two on 26.X.19XS. To give 
rhe tadpoles rough surfaces on which to attach, 
nlg.t-eovcred atones were inlioduced It.' the aqttatta 
when ihe spawn hatched, 

IX'IXirtmenr ol 7oolo^v, University ol Adelaide, C».P.O_ 
Box 498, Adelaide. S.*AuM. 5001 

Samples of embryos and larvae were collected aa 
shown in Tabic I and picsetved in Tyler's fixative 
(Tyler l%2), 

Measurements of developmental stages were 
taken using vernict callipers reading lo 0.05 nun. 
Illustrations were made wiih fhe aid of Wild MS 
stereodissccting microscope and camera lueida, 

Developmental slaves were clarified according 
to Oosner (I960). 


Ampk'vi/s: Ampk-xus was axillary (l-ig. I). The 
pair was ampleetani upon arrival in the laboratory 
possibly lu response to the reduced au pressure in 
the cargo hold ol the aeroplane (M. ,1- Tyler it M. 
Davics unpubl. obsT 

Eggs: Oviposition look place overnight, 2 and 
3.ixJ98&. Tempera! ure o\' the water in which the 
eggs were laid was 18,5°C. The egg mass cft'843 CfcjtS 
was 67.5 mm at its greatest width and compact 
Indisidual jelly capsules were tndnly adheient to 
cadi oilier (Fig. 2). The animal pole was dark brown, 
and approximately half of each cyg on ihe vegetal 
hemisphere was while. Only one membrane was 
detected in each capsule- 
Mean diameter of six eggs was 2.25 mm trance 
2.10-2.56 mm) and of the capsules, 2.98 mm (range 
2.60-3,60 mmk On the day following oviposition. 
rhe eggs were in tare cleavage (Sfa,ee 9). Within 2<t 
hours (hey had progressed to the neural fold siage 
(Stage 14) and within a limber 24 hours, the 
embryos were at Stage 17. Ihc cmbrvos wcrc 
peiched on a large choeolale-colouted yolk sac (Hit:,. 
3); large erescentic adhesive onj.ans were located 
anterodorsally, A prominent stotnodaca! pit was 
located posterodorsallv to the adhesive ornan (lie 

On 7.i\.l9KH, the embryos remained in (he jelly 
capsules. The adhesive organs wee beginning to 
divide as head flexion commenced. The siomodacai 
notch was prominent and Ihe pronephros wav 
apparent laterally; the uill plate w«s clearly 



delineated. The lail was differentiated and the 
embryos were at Stage 18. 

Larvae: The embryos harehed on 8,ix.l988 al 
Stage 19. The head was flexed ventrally and (he 
adhesive organs situated ventrolatcrally as two 
raised pigmented suckers. A sromodaeal depression 
was dorsomedial to the suckers. The primordia of 
two pairs of gill filaments were present and eye 
bulges were apparent {Fig. 4a). The pronephros was 

still apparent. A large yolk sac provided nutrition 
lot the embryo. 

By 9.i\.1989 the embryos were at Stage 20. The 
two pairs of gills U'cre now filamentous, the anterior 
pair with four to five branches, the posterior pair 
with one to three. Ventral suckers were ventrolateral 
to an open mouth. The optic bulges were well 
developed but opaque. Nutrition was supplied by 
a large yolk sac (Fig. 4b). 

TABLE I. DitDensions of developmental stages of Liioria eucnemis. 


(days, date) 


Body length 
\, range in 

parentheses (mm) 

Total lentil h 

\, range in 

parentheses (mill) 











(26.x. 1989) 

(26.x. 19891 

(26.x. 1989! 

(26a. 1989) 

(26.x. 1989) 




(7. xi. 1989) 


(7 xi.1989) 











{2.96 -3. 181 
































5 ^7 

















(26.(1 2*. 5) 







(8. 7-9.6) 



























11. 1 




29. fi 
















Fig. I. Amplcctant pair of Litona eutnemts. 

By I0.ix.t988, the larvae remained at Stage 20 but 
olfactory pits were now clearly visible. Larvae had 
reached Stage 21 within the nexl 24 hours. The 
cornea was transparent, pigmentation o^ the 
adhesive organs was fading and the gills were more 
filamentous (Fig. 4c). Ridges could be distinguished 
on the lateral and posterior margins of the mouth, 
Foreshadowing the labia] teeth and papillae. The tail 
had elongated; nutrition was still supplied by yolk. 

By 15. ix. 1988, the external gills were completely 
covered on both sides by the operculum. The 
adhesive organs were degerating, although some 
pigmentation remained. The horny beak was 
keratinized, but the tooth rows had not yet formed. 
Nutrition was still provided by yolk and the embryo 
was at Stage 24. Some tadpoles were now free 

iig. 2. Dorsal view of a spawn clump of l.iuma eucnemts. 
Greatest diameter is 67.5 mm. 

The spiracle had begun formation by 17.ix.1988. 
It was situated venlrolaterally and the opening 
directed posteriorly. Two upper and the two lower 
entire tooth rows were keratinized. The gut was 
loosely coiled but yolk-filled, and embryos were at 
Stage 25. 

Larvae were not sampled again until 27.ixJ988. 
The spiracle had elongated and its orifice was now 
directed posterodorsally. Mouthparts were com- 
plete, the larval gut had formed and the tadpoles 
were feeding on vegetable matter. The development 
oi' the limb buds indicate that the tadpole was at 
Stage 26. 

Fig. 3. A) Dorsal and B) anterior views of embryo at 1 imnu ei/aiemis at Stage 17, Scale bar represents 1 



Fig. 4. Litoria eucnemis larvae: A) lateral view at hatching, Stage 19; B) lateral view Stage 20; C) lateral view Stage 
21. Scale bar represents 1 mm. 

OF VliLOl'MJ.NT IN UfORfA £UC\r\!fS 


J«m SSffiflS (CT 1 Vl ™ " nd C) mQU ' h ** °' * b,Va * *^**MH8*f Sfcfcfi 31. $u|te bar repress 

The next sample oHarvae, taken on 5a. 1988, was 
at Stage 27. 1 he body of the larva was covered with 
brown pigment dorsally, and was pale ventrally. A. 
disrupted line of pigment granules bordered' the 
dorsal myotomes and fun her pigment granules svere 
scattered laterally and on the dorsaffin. 

Dimensions of developmental stages of L. 
cucnemis arc shown in (able 1. A larva ai Stage 31 
preserved on 26A.N88 is illustrated in Fig. 5a,b. The 
anus was dextral and the spiracle clongaie and 
sinistral. The dorsal fin was shallow and tapered 
to a gem ly -rounded tip, General body coloui was 
brown with faint pigment granules on (he tail 
myotomes and the dorsal fin. 

The mouth disc of a larva at Stage 31 is shown 
jji Fig. 5c. There are two upper and three lower rows 

of labial teeth. Both the second upper and the first 
lower row of teeth have a median gap. There are 
two tows of labial papillae lorming a v\ide band 
around the lateral and posterior margins of the 

Metamorphosis: Ry lO.xi 1988. the left \orearm 
had protruded in one larva and the firsr froglci ai 
Stage 46 was recorded on i6.xi.19K8, a total of 74 
days after the eggs were laid and 69 days aftci 
hatching. Water temperatures varied throughout the 
period of development from Jfi,4 -23.S a C 

Larval behaviour: Larvae frequently aggregated, 
mostly beneath the stones within I he aquaria or on 
the surface of the lettuce. 'I hey attached themselves 
by their suctorial mouihparts to the floor of the 
aquaria during water changing, 




I 'lie clcwlopjneul of Lito>'i(t nuwims is unusual 
amonasi Australian hyhds trt utilising a lai^e solk 
rescue and .showing relauveiv lute development ol 
the IftrURl gift l 1 * 11 -' species hatches at stage lil nm 
the jmt docs not fomi uniil stage 2$, 4-19 days alter 
hatching). Such development is more representative 
of direct -developing •vpecies or those t haj spend a 
considerable period of l hen development within the 
egg capsule [tup Kyarwrtft yihuitnuotus (Moore 
I%1; Ansli.v I9S1U- Amongst Ansiraiian species, 
this mode ol development is restricted to 
leptodaetylid frogs. /.. tSWCWU/S is u New Laiinea 
specie v that extends us range FiitO Australia, and its 
lelaiionships he with the new Gumcan hylid fauna, 
in particular the montane .speck's ol Lttvnti -and 
Nyctimysies. huch species lay laJgC unpignicrHcd 
eggs; tadpoles, where Known, exhibit louc 
adapiations (Men/ies 1974; Met' & Davies 1V7V: 
ZweHel 1983). However, complete life history daia 
are unknown for ihese species. The cyjis of L t 
eucficmis are latjic in comparison with olher 
Australian hylids (see lylci 198s) lor review), dirccily 
ivUed to the mode of early development. [Montane 
New Guinea i.itotat and \ ! yi'{itny\U\s species also 
are described as having laige eggs (Men/.ies l u 74)), 

The single adherent etfg mass is an adaption in 
IMonu QUCtwmis lo a lotic lifestyle: the tooth rows 
and oiher sti ueiurcs are typical of other Australian 
hvlids (Martin & Watson W7JJ, hut the suctorial 
mouth disc is an adaptation id fast-flowing waters, 

Amongst Auslralopapuan ami runs, there mt: 
several kinds of lot ic adaptations o) (he mouth disc. 
The extreme elaborations exhibited b> t.ttoriti 
m{tWofi\, I- rhi'fKtihi (Liem I97+)j /.. urlnkhiuM 
{Men/icx 1974», and Sycfhuystes tluy> (see l_vler 
Vffii lor Plate) consist of a large suctorial smiciurc 
occupying the entire ventral prcocular region of the 
tadpole, luuductylus spp have umhreJIa-.diaped lips 
hanging vcntroincdially (Liem &. Kosmer 1973) 
whilst in L. subyhtthiulo^i \ht tuouthpaits form a 
funnel in which all labial tooth rows and 
pigmentation of ihc homy beak are absent Oral 
disc tubercles have keratinized tips whilst unusual 
black filaments are suspended anteriorly to the. 
pharyns ( lylct & Anstis 1975). the Jess elaborate 
adaptations occupying a lesser proponion ol the 
onieiovcntral surface of ihe tadpole and exhibited 
by / wwrtemis aie shared by L cuphmdi and /.. 
tneirkwii (Tyler ct ai. 1983), L. testttvtri (Martin & 
Wahon 1971), Me&istolath Hyjtunus Myler « uL 
1979) and Misofthya, hulhus (Watson k Martin 

Other torreni adaptation* include the narrow (iiil 
fins and the Ventrolateral spiineular opening, 


1 thank Michael Mahony for the provision of the 
adult frogh- Michnd 1. lylet assisted with tadpole 
rearing and crilicalh ivad the manuscript. This 
work was supported by University of Adelaide 
Department o\' /oology funds. 


Ansiis VI. <WM| Wacduie Ixalogv and rawyc on-ii-aon 
(Ml lire \cv 'mmiiIi Wale hoy, h.\uitituu>; ^hu^tiiohis 
tAnuf.i: J cpioduetvhiJ.ici. Aim I HPQ&Ml I. 1-9. 

CiKtfiEU H (•". (Iwfi) 'RcimiIl's and Aijinhihiuns ol 
Australia" ?rd edin (Keed, Swhtey). 

{ o s\i ft, K* IWli A simpliiicd tabU' lo. sraginu anuian 
(•uilnvos .md huu\y wirh notes oi> .d^niihcaiion. 
Hcri^iohnui-ti 10. IHV-IVtl. 

tllM, I). S. (19741 \ r.'vu-W of tllC Ufentt rum/ioli': 
sno;ic- gioiip and a dfseopnon of u fltf^ spv; k v - i^l 
/ itoritt fi"ih nonht*ro Oucenvliind, Australia ^iuuh. 

nvhdaci. M&n CW.M«y. 17(1). im-16s. 

(t Ho-.mlh. VV. [Ml\) Ffftjit« «u ilu- Kemis 

Tiituhii't\lus Nsiih ifcHTjpitno; "' ,w,> Mew spectcs 

lAnuia* U'l'lodjaylidiic}. Wfc/M \ki Mm W^\ 

4«s ia?, 
MvktK, A. a. (fc W,\r,ns, (i. f. (1971) Life hisiin y ;»s nn 

aid ty gciKTK- delimiialion in tlie launlv 1 1\ tidac. u#ij 

(.1971,, ?S ?9, 
Mi \/,is I. 1. (197411 H.mJbook ol\'Miiti»iun Nft» Guinea 

f«M!V. WAti I coloyy 1n\iiint^ Handbook no I. 
\ir,..Hi I A. (I%1) 1 -io"s ot ea>lcin New Somh Walci. 

flvfi-Atk 'Wrt tttfi //oV. 121, l49--^fs 
Tmtk M. f (tW) On the proscmitior. oi amnan 

uUnotw ,4;o/ ( 7. s'ci. 25, 222; 

_ (»WiS> Papuan hylid frogs ol nw genu:- //v/« it^wA 
fftfutrirJ. l.vuten %, \-Z\KK 

"Aiisnuliriii liwt" I Vikinj: <>'>» 


A Avhis, M. (I^;S) Taxoiioiny mid biolouy ol 

I'rtJga ol ihu / Htwut t/zw/weomples (Atianr Hvhclji;), 
AVr, .S*. ,1ro7. U/o. P, 4I-.S0. 

Crouk <i. A. &Davil> M <I9K'JJ Ueproducfivc 

hiolouv oi iIk 1 I i* >v.' s of the Ma(?,C*fl Vuvk •iysifin. 
NoTihern Tcnilu)> thtrtAH, 41^ 444), 

& Dwils. M. (1979) kiHletiniiion and lilV 

vvoluiionni'v jrtrfti of (he AiNiMlop.-i|-iuanj'rog Ktiifus 

Nyvtwty*f>:)> Nieiiuw. t(M/. ./. 2Vo^, 27, "■>5-77,:, 
, VI.MU.N. A, A & l")wil,x, M rt^rftj t'iiolniiv.tind 

svsieiiiaii'.s ol '< ni*^ Iminotlvnivaine jjenus tAnnui; 

U-pioda.ivlidac) from uoillmeMcni MiMmlia. Aust. I 

/.<H)L 27/135-1^0. 

A Waison, O. K OW-jOn i he nomcnelainrc o.' 

a tiytiil nvc froii tVom Ot'^ndanJ, hvns. i<. Sue. .S. 

,4fr5f WWW), l 1 ^ N4. 
Wmson. O. h ft MA«1 in '\. A. i!973) Lil'c Insnn v, l.irval 

morpholoev and ielrttionshi|v. ol' MiMraliau le(Ho- 

dacivlid IrcagJw /toct B, - i - 1 -45. 
/.vviuii. R. G. iiy^O) Kesidis ol 

IV|iediiior^ no Kit Ito^ and (i/ani* 

IVmitvnlo. Kipua New Ouinctt Aifl 

//r.5/. 165(^). W0-4J4. 
(1991) Tu(> nc'vv livlid l'(ow% liom I'apnit New 

iiiitiKM in*) .i discusMo'n ol il»« : \\fttM\'W\ f&tfwa 

speen-^ ; MOii|S- Am* Mitt. UviittiK* -75^. 1 1\. 

the Aiclit'Ol.1 
I'roin the Uuon 
,l///, \-tus. HU(, 



by Gary R. Beardsell 


ftmeettoro <>/ ,hr Hojful Kockrty ■■■/ s. test* (1989), H3(4t h\ JJ4 



A study of intraspeedie variation among the rhnee 
disjunct populations 01* Litoha chloris (Boulengerl of 
Eastern Australia showed thai the northern population 
differed in morphology from Ihe other two populations'. 

Isoenzyme electrophoresis demonstrated extensive 
genetic divergence between the northern and southern 
populations resulting |Q (he recognition of the northern 
population as y distinct species, L .\antfiorrtera\ 

Early development and larval moiphology of / . chloris 
have been described ' and Ihe laivae of /.. xanthonnru are 
identical to those of /.. chhris'. 

Merc I genetic compatibility through a fertile cross 
between a captive male L. vhlons and a female L. 

two adult male /_. chhtis were acquired from Finch 
I Utton Gorge, MaCkay, Queensland in lamiary 1984, and 
one adult female of L xanthome™ from Crystal rails near 
Cj.ii ms. QIJ on J0.xii.1987. The frogs were housed lf| in 
indoor tenaiium. Amplexus occurred on 28.1.1988. 

Embryos were reared m an aquarium with rainwater to 
a depth of 24 cm. VVatei icmperature was IX-29°C. Larvae 
Were fed pond algae, commercial fish food and boiled 
lettuce. When they approached metamorphosis ihey were 
transferred to a icrrarium. The froglOU Wefe fed on house 
flics and flour moths. 

Measurements were made using vernier calipers reading 
to 1 mm and an ocular mietomelcr to 0.1 mm. The larval 
mouth disc was drawn under a stereoscopic microscope 
while the larval and froglet drawings were inked directly 
onio enlarged photographs, that were later treated with 
potassium iodide and a sodium thiosiilphate living bath 
to remove Ihe photographic image and iodine stain 
icspcctively 1 . Developmental stages follow GosneC 

Calling commenced on l7.xi.J987 aui _\ eontinutd on 
most nights through to mid-January when the female L. 
xanthamcra became gravid. Amplexus occurred on 


►Ig. I. Developmental stages Of a iertile cross between 
Litoriachlom and A, xamhomera. A, Stage 29; B, Stage 
i4; r, stage 42. Scale bar - I mm. 

28.ii.J988 at 0800 hr, and the male continued to call ffli 
one minute while the female produced a soil call Hie 
• iitiplectanl pair was then placed Into a smaller breeding 

Oviposiiion OOCUrted from 0630 to 0700 hr on 29.ii.l988. 
Approximately 600 eggs were laid as a Bat, jelly-like mass 
on the surface of the water; mean ovidiameter =* 1.5 mm, 
capsule diameter - 4.4 mm, in = 6). I he animal pole 
Was dark brown, the vegetal white. The fertile eggs began 
to sink and adhere to the bottom and sides of the 
aquarium in ihe first hour. 

Dimensions of the embryos and larvae are shown in 
Table I. Hatching began l.iii.1988 at stages 20 & 21, The 
embryos were dark to mid brown; length 6,5 mm. Each 
embryo had three pairs of external gills with numerous 

On 3,iii.l988 the larvae were at stage 23, The operculum 
developed and melanophores appeared along the mni- 
dorsal tine, over ihe tail musculature and as circular 
patches over the dorsal surface of the abdomen. The larvae 
were pale brown and the coils of the intestines visible 
through the abdominal wall. The larvae i cached stage 25 
ai 1700 hi (4.iii.l988). 

At stage 29 (Hg. )A) the larvae had a total length ot 
15.5 mm and a body length of 5 mm, Most laivae 
developed a lateral curvature of the tail o\ varying degrees 
of severity between stages 28-30 in weeks 4 & 5 Ai this 
point there was marked variation in growth: some larvae 
attained :i total length 2x that of others. As the larvae 
approached stages 34-38 between weeks 5 and 7 the 
severity of the tail bend facreafied into an S-shape in many 
individuals (h'ig. 2). At stage 34 body colour was light 
brown with a traiispaiem area over the abdominal wall. 
Pigmentation was apparent on the dorsal tail fin and tail 

seiilature (Fig. (H). On day 49 the largest larva 

nicM-iirei! 52 mm in stage 40 (I ip. tC ) 

1 He month disc (fig. 3) of a larva in stage 34 | ( ;»s the 
typical hvlid formula of 

I I 

|ig. 2. Abnormal tail bend in a i.vt...o* larva at Stnjjr 40 
Total length 52 mm. 


TABLE 1. Dimensions of developmental stages of a cross between Litoria chloris and L. xanthomera. 

Means (mm) with ranges in parentheses 


(Hours) (N) 

1.5 4.5 

(1.5-1.6) (43-4.7) 


















10 & 11 


































































In many larvae extra papillae with labial teeth appeared brown with a very fine, diffuse speckling of white 

between the upper rows of labial teeth (Fig. 3). disappearing after one week. 

Four froglets completed metamorphosis 20.iv.1988 and 
had snout-vent (S-V) lengths of 14.4, 14.6, 14.8 and 15.1 The embryo and larval lifespan extended over a period 
mm. The remaining 52 surviving larvae completed of 51-56 days at 18-29°C. The froglets were measured a! 
metamorphosis by 26.iv.1988. The majority of juveniles monthly intervals after completion of metamorphosis (Fig. 
metamorphosed with pelvic girdle abnormalities (Fig. 4A) 5). Within a few days they developed a white supralabial 
to varying degrees of severity. Approximately on third of stripe disrupted in some individuals and complete in 
the juveniles displayed no external sign of skeletal others. The mid-brown colour changed to a light mustard- 
abnormalities (Fig. 4B). The colour of juveniles was mid- brown within two weeks of metamorphosis. By 5.V.1988 


Fig. 3. Hybrid larval mouth disc al Stage 34. Scale bar 

=■ 1 mm. 

Fig. 4. A, Pelvic girdle abnormalities in a three month old 
hybrid froglct (S-V 23 mm); B, Norma! three month old 
hybrid froglet (S-V 27.2 mm). 

the supralabial stripe developed a green tinge while green 
pigmentation developed on the supraobital skin. 

The thighs developed the orange of L. xanthome™ by 
May-mid June, the supralabial stripe became indistinct 
concomitantly, with a green tinge over the brown on the 
body parts that are green in the adult life. After eight 
months all 25 surviving froglets continue to grow 
regardless of the presence or absence of pelvic girdle 
abnormalities. On 6.xii.l988, the largest froglct had a 
snout-vent length of 37 mm and had developed nuptial 

The life history pattern of the hybrid conforms to the 
pattern of Australian hylids". 

The larval life span of 51-56 days is in agreement with 
the 41 days recorded for L. chloris { and 58 days for L, 
infrafrenuta considered usual among tropical species 
adapted to life m ephemeral waters'*. 

More importantly, development of a lateral curvature 
of the tail in larvae at four weeks manifesting as pelvic 
girdle abnormalities in newly metamorphosed froglets 
supports the premise of genetic differentiation attained 
by L. xanthomera and its recognition as a separate taxon 
from L. chloris 2 . The development of some apparently 
normal froglets suggests that these species are closely 



t» 25- 


• 20- 






I I 

1- n-« 



12 3 4 5ft 

71m« from Metamorphosis (months) 

Fig. 5. Growth rate of hybrid offspring from a cross 
between Litoria Moris and /.. xanthomera. (Mean and 
range; n = sample size). 

I thank Mr M. J. Tyler and Dr M. Davies who critically 
read the manuscript. Special thanks are due to Richard 
Perkins for his expert skills in photography and 
introducing me to the technique of bleach-outs and the 
science department of Ingle Farm High School for the 
use of various equipment and their general interest in the 
frogs, Thanks are due to my brother Kym for typing the 


Davies, M & McDonald, K. R. (1979) Aust Zooi 20, 

2 & Adams, M. (1986) Proc. R. Soc. Vict. 98(2), 

'Watson, G. F. & Martin, A. A. (1979) Aust. Zooi 20, 

4 Palerson (1979) "Book of the Darkroom". (Paterson 

Products Ltd. Sawtells of Sherborne Ltd, England.) 

Gosner, K. L. (1960) Herpeto/ogica 16, 183-190. 
''Martin, A. A. & Watson, G. F. (1971) Copeia (1971) 

Banks, C B.; Birkett, J. R.; Dunn, R. W. & Martin, A. 

A. (1983) Trans. R. Soc. 5. Aust. 107(4), 197-200. 
"Duellman, W. E. & Trueb, L., (1986) Larvae, Chapter 6 
in "Biology of Amphibians". (McGraw-Hill Book Co., 
New York.) 

GARYR. BEARDSELL, Ingle Farm High School, Belalie Road, Ingle Farm, S. Aust. 5098. 





G. M. E. MAYO, B.Ag.Sc, Ph.D., D.Univ. 


C. J. M. GLOVER, J.P., M.Sc 

N. A. LOCKET, M.A., B.M., B.Ch. f Ph.D., D.O. 

G. A. CROOK, B.Sc. 

R. H. WATERS, J.P., F.A.LM, 






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